Proposed Deepwater Lophelia Coral HAPCs Page 1 of 7
Proposed Deepwater Lophelia Coral HAPCs
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Identification_Information: Citation: Citation_Information: Originator: Florida Fish and Wildlife Conservation Commission (FWC), Fish and Wildlife Research Institute (FWRI) Publication_Date: January 2008 Title: Proposed Deepwater Lophelia Coral HAPCs Geospatial_Data_Presentation_Form: vector digital data Other_Citation_Details: The following reports and published manuscripts were used to identify or describe deepwater coral habitat in the South Atlantic Fishery Management Council's jurisdiction. Overview and Summary of Recommendations: Joint Meeting of the Habitat Advisory Panel and Coral Advisory Panel 2004, 2006, and 2007; General Description of Distribution, Habitat, and and Associated Fauna of Deep Water Coral Reefs on the North Carolina Contental Slope (Ross, 2004); Deep-water Coral Reefs of Florida, Georgia and South Carolina: A Summary of the Distribution, Habitat, and Associated Fauna (Reed, 2004), Habitat and Fauna of Deep-Water Coral Reefs off the Southeastern USA - Report to the South Atlantic Fishery Management Council Addendum to 2004 Report 2005-2006 Update- East Florida Reefs (Reed, 2006), and Review of Distribution, Habitats, and Associated Fauna of Deep Water Coral Reefs on the Southeastern United States Continental Slope (North Carolina to Cape Canaveral) (Ross, 2006). Online_Linkage:
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Amendment currently under development.
At their joint meeting in Charleston in November 2007 Habitat and Coral Advisory Panels received recent research updates on on the status and distribution of deepwater coral systems in the region. Based on this new information, the Panels proposed to reaffirm the 4 areas and add a fifth, the Blake Ridge Diapir. The Council subsequently voted to adopt the Panels' proposal for inclusion into the public hearing draft for the Comprehensive Ecosystem Amendment. Purpose: The spatial representation of proposed Coral Habitat Area of Particular Concern C-HAPC areas in the South Atlantic Region. Supplemental_Information: Prior to July 1, 2004, the Fish and Wildlife Research Institute (FWRI) was known as the Florida Marine Research Institute (FMRI). The institute name has not been changed in historical data sets or references to work completed by the Florida Marine Research Institute. The institute name has been changed in references to ongoing research, new research, and contact information. Time_Period_of_Content: Time_Period_Information: Single_Date/Time: Calendar_Date: 2007 Currentness_Reference: publication date Status: Progress: Complete Maintenance_and_Update_Frequency: As needed Spatial_Domain: Bounding_Coordinates: West_Bounding_Coordinate: -81.131245 East_Bounding_Coordinate: -75.690186 North_Bounding_Coordinate: 34.410302 South_Bounding_Coordinate: 24.182720 Keywords: Theme: Theme_Keyword: Benthic Habitats Theme_Keyword: Corals Theme_Keyword: Lophelia Place: Place_Keyword: South Atlantic Bight Access_Constraints: Available without restriction Use_Constraints: FWC-FWRI must be credited. This is not a survey data set and should not be utilized as such. These data are not to be used for navigation. Point_of_Contact: Contact_Information: Contact_Person_Primary: Contact_Person: Tina Udouj Contact_Organization: FWC-FWRI (Florida Fish and Wildlife Conservation Commission-Fish and Wildlife Research Institute) Contact_Position: Assistant Research Scientist Contact_Address: Address_Type: mailing and physical address Address: Fish and Wildlife Research Institute 100 Eighth Avenue Southeast City: St. Petersburg State_or_Province: Florida Postal_Code: 33701 Country: USA
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Contact_Voice_Telephone: 727-896-8626 Contact_Facsimile_Telephone: 727-893-1679 Contact_Electronic_Mail_Address: [email protected] Hours_of_Service: 8:00 a.m.-5:00 p.m. Eastern time Security_Information: Security_Classification_System: FWRI-DC Security_Classification: Available without restriction Security_Handling_Description: Available without restriction Native_Data_Set_Environment: Microsoft Windows XP Version 5.1 (Build 2600) Service Pack 2; ESRI ArcCatalog 9.1.0.780
Data_Quality_Information: Attribute_Accuracy: Attribute_Accuracy_Report: Feature attributes were visually inspected and appear to be correct. Logical_Consistency_Report: No logical inconsistencies are known to exist in this data set. Completeness_Report: All HAPC areas defined as of January, 2008 are represented. Lineage: Source_Information: Source_Citation: Citation_Information: Originator: NOAA Coastal Services Center Publication_Date: 20070725 Title: Southeast Bathymetry Geospatial_Data_Presentation_Form: vector digital data Other_Citation_Details: Southeast regional bathymetric contours consist of a vector coverage of bathymetric contours with increasing resolution in coastal areas. Contours were derived from a composite of several bathymetric datasets of varying regional coverage and resolution. Isobath intervals range from 2 meter in coastal areas to 200 meters in deep offshore areas. Online_Linkage:
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Source_Citation_Abbreviation: NCDDC Bathmetry Source_Contribution: The 300 meter and 400 meter isobaths were used to generate portions of the Stetson/Savannah and East Florida Lithoherms/Miami Terrace CHAPC boundary. Source_Information: Source_Citation: Citation_Information: Originator: Skidaway Institute of Oceanography Originator: Clark Alexander Originator: Mike Robinson Publication_Date: 20040915 Title: Coral Mounds Geospatial_Data_Presentation_Form: vector digital data Other_Citation_Details: Coral Mounds were digitized from the Bottom Character Map of the Northern Blake Plateau OFR-93-724. For more information see:
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Process_Description: Dataset modified based on recommendations of the 2007 SAFMC Joint Meeting of the Habitat and Coral Advisory Panels. The C-HAPC surrounding the Blake Ridge Diapir methane seep live-bottom area was appended to this data set. Process_Date: 2008
Spatial_Data_Organization_Information: Direct_Spatial_Reference_Method: Vector Point_and_Vector_Object_Information: SDTS_Terms_Description: SDTS_Point_and_Vector_Object_Type: G-polygon Point_and_Vector_Object_Count: 5
Spatial_Reference_Information: Horizontal_Coordinate_System_Definition: Geographic: Latitude_Resolution: 0.000000 Longitude_Resolution: 0.000000 Geographic_Coordinate_Units: Decimal degrees Geodetic_Model: Horizontal_Datum_Name: North American Datum of 1983 Ellipsoid_Name: Geodetic Reference System 80 Semi-major_Axis: 6378137.000000 Denominator_of_Flattening_Ratio: 298.257222
Entity_and_Attribute_Information: Detailed_Description: Entity_Type: Entity_Type_Label: prop_dc_0607_area Attribute: Attribute_Label: FID Attribute_Definition: Internal feature number. Attribute_Definition_Source: ESRI Attribute_Domain_Values: Unrepresentable_Domain: Sequential unique whole numbers that are automatically generated. Attribute: Attribute_Label: Shape Attribute_Definition: Feature geometry. Attribute_Definition_Source: ESRI Attribute_Domain_Values: Unrepresentable_Domain: Coordinates defining the features. Attribute: Attribute_Label: NAME Attribute_Definition: Name of proposed Coral-Habitat Area of Particular Concern (C-HAPC). Attribute: Attribute_Label: AREA_METR2 Attribute_Definition: Area in square meters. Attribute: Attribute_Label: AREA_MILE2 Attribute_Definition: Area in square miles.
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Distribution_Information: Distributor: Contact_Information: Contact_Person_Primary: Contact_Person: GIS Data Librarian Contact_Organization: FWC-FWRI (Florida Fish and Wildlife Conservation Commission-Fish and Wildlife Research Institute) Contact_Position: GIS Data Librarian Contact_Address: Address_Type: mailing and physical address Address: Fish and Wildlife Research Institute 100 Eighth Avenue Southeast City: St. Petersburg State_or_Province: Florida Postal_Code: 33701 Country: USA Contact_Voice_Telephone: 727-896-8626 Contact_Facsimile_Telephone: 727-893-1679 Contact_Electronic_Mail_Address: [email protected] Hours_of_Service: 8:00 a.m.-5:00 p.m. Eastern time Distribution_Liability: This data set is in the public domain, and the recipient may not assert any proprietary rights thereto nor represent it to anyone as other than a FWC-FWRI produced data set; it is provided "as-is" without warranty of any kind, including, but not limited to, the implied warranties of merchantability and fitness for a particular purpose. The user assumes all responsibility for the accuracy and suitability of this data set for a specific application. In no event will the staff of the Fish and Wildlife Research Institute be liable for any damages, including lost profits, lost savings, or other incidental or consequential damages arising from the use of or the inability to use this data set. Standard_Order_Process: Digital_Form: Digital_Transfer_Information: Format_Name: shp Transfer_Size: 0.003 Fees: None. However, persons or organizations requesting information must provide transfer media (CD- ROM only) if FTP is not available and must pay express shipping costs if express shipping is required. Ordering_Instructions: Contact GIS Librarian by e-mail, telephone, or letter explaining which products are needed and providing a brief description of how the products will be used. Also, provide name and address of the person or organization requesting the products. Turnaround: Usually within 10 business days, although, complex requests may take longer Custom_Order_Process: Contact GIS Librarian
Metadata_Reference_Information: Metadata_Date: 20080208 Metadata_Contact: Contact_Information: Contact_Person_Primary: Contact_Person: GIS Data Librarian Contact_Organization:
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FWC-FWRI (Florida Fish and Wildlife Conservation Commission-Fish and Wildlife Research Institute) Contact_Position: GIS Data Librarian Contact_Address: Address_Type: mailing and physical address Address: Fish and Wildlife Research Institute 100 Eighth Avenue Southeast City: St. Petersburg State_or_Province: Florida Postal_Code: 33701 Country: USA Contact_Voice_Telephone: 727-896-8626 Contact_Facsimile_Telephone: 727-893-1679 Contact_Electronic_Mail_Address: [email protected] Hours_of_Service: 8:00 a.m.-5:00 p.m. Eastern time Metadata_Standard_Name: FGDC Content Standards for Digital Geospatial Metadata Metadata_Standard_Version: FGDC-STD-001-1998 Metadata_Time_Convention: local time Metadata_Access_Constraints: No restrictions on metadata Metadata_Use_Constraints: Metadata must be distributed with the data set. Metadata_Security_Information: Metadata_Security_Classification_System: FWRI-MC Metadata_Security_Classification: Available Metadata_Security_Handling_Description: Metadata must be distributed with the data set. Metadata_Extensions: Online_Linkage:
Generated by mp version 2.8.6 on Fri Feb 08 08:04:25 2008
file://C:\Documents and Settings\roger.pugliese\My Documents\proposed deepwater lophe... 2/19/2008 SOUTH ATLANTIC FISHERY MANAGEMENT COUNCIL
4055 FABER PLACE DRIVE, SUITE 201 NORTH CHARLESTON, SOUTH CAROLINA 29405 TEL 843/571-4366 FAX 843/769-4520 Toll Free 1-866-SAFMC-10 email: [email protected] web page: www.safmc.net
George Geiger, Chairman Robert K. Mahood, Executive Director Duane Harris, Vice Chairman Gregg T. Waugh, Deputy Executive Director
JOINT MEETING OF THE HABITAT AND ENVIRONMENTAL PROTECTION ADVISORY PANEL AND CORAL ADVISORY PANEL (November 7-8, 2007)
The Charleston Marriott Hotel 170 Lockwood Boulevard, Charleston, SC 29403
JOINT HABITAT AND CORAL AP FINDINGS AND RECOMMENDATIONS
Issues addressed and presentations made at this meeting included: 1) The Complexity of Habitats in the Proposed C-HAPCs presented by John Reed, Harbor Branch Oceanographic Institution; 2) Research Cruise Updates; Understanding Species use of Deepwater Habitats presented by Steve Ross, UNCW; 3) Cooperative Multi-Platform Mapping Cruise June 07 presented by Greg McFall Gray’s Reef National Marine Sanctuary (GRNMS); 4) Update on Mapping and Characterization on the Charleston Bump and Blake Plateau presented by George Sedberry, Superintendent, GRNMS; 5) Development of Rapid Assessment Tool (SEADESC) and Integration into Habitat and Ecosystem IMS presented by Steve Ross/UNCW and Tina Udouj/FWRI ; 6) ESDIM Deepwater Habitat Completion Report presented by Tina Udouj; 7) Development of Deepwater Habitat GIS and Habitat and Ecosystem IMS in support of Deepwater C- HAPC designation presented by Tina Udouj; 8) Panel Recommendations on Proposed Deepwater Coral HAPCs for the Comprehensive Ecosystem Amendment; 9) Member Comments and Recommendations for Enhancing, Expanding and Refining the Fishery Ecosystem Plan for the South Atlantic Region; 10) State Sub-Panel Breakout Sessions – Priority State and EFH Conservation Recommendations; 11) Development of Liquified Natural Gas (LNG) Facility, EIS and Larval Fish Research and Ocean Turbine Development presented by Jocelyn Karazsia, NOAA Fisheries; 12) in the SA Region presented by Jocelyn Karazsia, NOOA Fisheries; 13) MMS Alternative Energy Report and Workshop presented by Roger Pugliese; 14) Energy Policy Revision; and 15) Research Area Development Process for Grays Reef National Marine Sanctuary presented by George Sedberry/GRNMS.
Joint Habitat and Environmental Protection and Coral Advisory Panels - Recommendations, November 8, 2007
Proposed Deepwater Coral HAPCs:
• The Panel strongly supports the designation of the entire C-HAPC as proposed below (Figure 1.) as supported by the best available science, and does not support designating portions of the whole.
• The proposed deepwater coral HAPCs (C-HAPCs) should be adopted and implemented as soon as possible. Growing pressure for new and more intensive uses of the EEZ, including potential energy development, mariculture and emerging deepwater fisheries, requires rapid designation. The Panels recommend shifting other measures that may delay C-HAPC implementation into FEP Comprehensive Amendment 2.
• New information compiled for the Council and presented to the Panels by John Reed and Steve Ross and others constitutes the best available science, and continues to strengthen the case for protection of this world-class deepwater coral ecosystem. Additional important coral features and related habitats continue to be found within the areas previously identified, including low relief features, ridges, sinkholes, pinnacles and escarpments up to 500’ tall (Figure 2).
• Research suggests potential for additional deepwater coral habitats in the following areas (Figure 3): 1) Some distance north of the current boundary of the Stetson/Savannah/Miami Complex, 2) Between the Miami Terrace and the Pourtales Terrace, and 3) to the southwest of the Pourtales Terrace. The Panel recommends additional characterization work on these sites, to be factored into future habitat protection amendments.
• Only one chemosynthetic (“methane seep”) live-bottom community has so far been documented in the US South Atlantic EEZ, northeast of the boundary of the Stetson/Savannah/Miami Complex, on the Blake Ridge Diapir. It should be protected as a separate C-HAPC.
• Τhe Panels reiterate the previous request to the Council to interact with the US and Bahamian governments to find ways to collaborate on research as well as protection measures for shared deepwater coral ecosystems. The Council could communicate with the Bahamian government directly through the U.S. Departments of Commerce and Department of State.
The Panels recommend the C-HAPC as previously configured (Figure 1).
Recommend establishing a C-HAPC (2 square mile) surrounding the Blake Ridge Diapir methane seep live-bottom area.
2 Joint Habitat and Environmental Protection and Coral Advisory Panels - Recommendations, November 8, 2007
Regulations in the Proposed Deepwater Coral HAPCs:
The Panels recommend that the following management actions be taken inside the C-HAPCs: Recommended management measures in all the deepwater coral HAPC sites include the following: 1) compile, characterize and track threats to deepwater coral ecosystems in the region; 2) as far as possible, limit damage from both fishing and non-fishing threats, using all available administrative tools; 3) prohibit all bottom-disturbing fishing gears;4) prohibit harvest of coral, coral reefs and live/hard bottom organisms(all taxa, including gorgonians and other soft corals) except as allowed through appropriately protective research protocols and procedures; 5) prohibit anchoring, grapples and chain, and 6) fully implement the Council’s deepwater coral research and evaluation plans.
The intent of these recommendations is to eliminate any commercial harvest that might be presently permitted under the coral plan in any deepwater coral HAPC, but to allow controlled collection for research purposes consistent with the Council’s Deepwater Coral Research Plan (i.e. as allowed by the Secretary). In addition, more work is needed to characterize potential damage associated with other bottom-impinging gears (e.g. damage that might occur with the use of weighted long-lines, planers and cannonball weights). The Council should clarify the types of allowable fishing activities, including transit through the C-HAPCs. Non-fishing impacts would be fully covered in the Fishery Ecosystem Plan and in future habitat policy statements.
Resources for Implementation
Despite the growing evidence that the deepwater coral ecosystems of the region constitute a world-class resource under rising threat, funds continue to be scarce for all aspects of management of these natural treasures. The Panels recommend that all possible sources be explored to obtain necessary funding for research and monitoring, outreach and education, and enforcement, once the C-HAPCs are emplaced. Funding must be found for high resolution mapping especially in areas of high probability.
JOINT HABITAT AND CORAL AP CONCENSUS RECOMMENDATIONS: Panel members were requested to provide comments on the potential list of actions for consideration in a developing Comprehensive Ecosystem Amendment. Discussions revolved around the existing list of proposed measures for the Amendment and the following recommended modifications are: Comprehensive Ecosystem Amendment 1 measures should: • Establish and protect expanded deepwater Coral HAPCs;
Comprehensive Ecosystem Amendment 2 measures should: • Establish a zero harvest for Sargassum; • Address octocorals harvest and quota level while considering octocorals as EFH; • Establish provisions to allow for the discovery of new octocorals species and new compounds (biomedical products), but not for mass exploitation and harvesting of species;
3 Joint Habitat and Environmental Protection and Coral Advisory Panels - Recommendations, November 8, 2007
• Consider invasive species highlighting lionfish in FEP and proposed or future Ecosystem Amendment; and • Establish Allowable Gear Areas (deepwater trawls and other gears as data allow)
Figure 1. Proposed Deepwater Coral HAPCs “South Atlantic Deep Reef Complex*” *New name proposed by Clark Alexander, Coral AP Member
4 Joint Habitat and Environmental Protection and Coral Advisory Panels - Recommendations, November 8, 2007
Figure 2. Deepwater Habitat associated with the proposed “South Atlantic Deep Reef Complex”.
5 Joint Habitat and Environmental Protection and Coral Advisory Panels - Recommendations, November 8, 2007
Figure 3. Areas Recommended for Future Research and Mapping of Potential Deepwater Coral Habitat.
6
SOUTH ATLANTIC FISHERY MANAGEMENT COUNCIL
ONE SOUTHPARK CIRCLE, SUITE 306
CHARLESTON, SOUTH CAROLINA 29407-4699
TEL 843/571-4366 or FAX 843/769-4520 Toll Free 1-866/SAFMC-10 E-mail: [email protected]
Web site: www.safmc.net
Louis Daniel, Chairman Robert K. Mahood, Executive Director George Geiger, Vice-Chairman Gregg T. Waugh, Deputy Executive Director
Overview and Summary of Recommendations
JOINT MEETING OF THE HABITAT ADVISORY PANEL AND CORAL ADVISORY PANEL
June 7-9, 2006
Wyndham Grand Bay Hotel 2669 South Bayshore Drive Coconut Grove, Florida 33133
Issues addressed at this meeting included: 1) Habitat and Ecosystem Webpage and Internet Mapping System; 2) Deepwater Coral Habitat Research and Management including the Development of a Deepwater Coral Research and Monitoring Plan for the South Atlantic Region; 3) Sargassum Research; 4) Listing of Elkhorn and Staghorn Corals as Threatened under ESA; 5) Fishery Ecosystem Plan and Comprehensive Ecosystem Amendment Development; 6) Snapper Grouper Amendment 14 - Deepwater Snapper Grouper Marine Protected Areas; 7) Summary of Updated SAFMC Energy Policy Statement; 8) Liquified Natural Gas (LNG) Pipeline Development: Assessing Impacts on Nearshore and Deepwater Coral Habitats; 9) Research Associated with Proposed LNG Pipeline Development; 10) Windfarm Development in the South Atlantic Region; 11) Initiation of the Development of a SAFMC Aquaculture Policy; 12) Invasive Species: Lionfish Research and Proposed Workshop; 13) Development and Management of Regional Ocean Observing Systems; and 14) National Habitat Plan and Southeast Aquatic Resources Partnership
- 1 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006
1) Habitat and Ecosystem Webpage and Internet Mapping System Roger Pugliese introduced the development of tools to support the move to ecosystem management included the Habitat and Ecosystem Internet Mapping Server and the Habitat and Ecosystem Section of the webpage. Tina Udouj of the Florida Fish and Wildlife Research Institute (FWRI) presented a summary of the development of the Habitat and Ecosystem webpage and Internet Mapping Server (IMS). Myra Brouwer conducted a live presentation detailing information presented and process involved in accessing the Habitat and Ecosystem section of the webpage. Roger Pugliese demonstrated access and use of the Internet Map Server. Panel members were requested to provide comments and recommendations that will aid in the further refinement of the Ecosystem site and IMS to better support regional ecosystem management.
Additional Background: The South Atlantic Council and the Florida Fish and Wildlife Research Institute (FWRI) partnered to develop a Comprehensive Habitat and Ecosystem webpage that is accessible from the South Atlantic Council’s web site. FWRI is hosting an Internet Map Server (IMS) application with links to downloadable bottom type data, associated metadata, substantial program information for the Council and links to related sites. The sit was transitioned to a web portal and is now operated and maintained through contracts with Mapwise Inc. and accessible and updated by Council staff. The Internet Map Server (IMS) component of this project brings the power of Geographic Information Systems (GIS) technology and Image Analysis tools to ordinary Internet browsers. The IMS will be an effective tool for displaying, sharing and querying coral and benthic habitat data and other pertinent ecosystem information across the South Atlantic region. In addition, researchers have a unique opportunity to access video and still imagery archives of coral and benthic habitats served from this site.
JOINT HABITAT AND CORAL AP RECOMMENDATIONS • Add metadata records; • Develop a mechanism for adding new data; • Add available water quality information; • Provide permission to access detailed data and get the latitude and longitude; • Expand data inland to include watershed and estuarine data; • Incorporate available LIDAR data including states’ data are from the estuary and within three miles of shore; • Include 35 years of North Carolina data that is now all digitized; • Add or link to Dade County data; and • Investigate adding 30-year water quality database for Biscayne Bay, which is not web-accessible.
2) Deepwater Coral Habitat Research and Protection In December 2004 the Council approved management actions proposed by the Habitat and Coral Advisory Panels to establish new deepwater coral HAPCs for inclusion into the Comprehensive Ecosystem Amendment.
- 2 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006
2004 Reports to Council and Advisory Panels- Dr. Steve Ross of the University of North Carolina at Wilmington (UNCW) and John Reed of the Harbor Branch Oceanographic Institute (HBOI) made presentations on deepwater coral distribution and characterization in the South Atlantic Region. Andy Shepard, Director of the UNCW/NURC was contracted to coordinate the preparation of the reports for the Council. The presentations encompassed exploration and characterization conducted to date on deep water coral habitats in the South Atlantic region. The following reports developed for the Council summarize this information: GENERAL DESCRIPTION OF DISTRIBUTION, HABITAT, AND ASSOCIATED FAUNA OF DEEP WATER CORAL REEFS ON THE NORTH CAROLINA CONTINENTAL SLOPE (Ross, 2004); and DEEP-WATER CORAL REEFS OF FLORIDA, GEORGIA AND SOUTH CAROLINA: A SUMMARY OF THE DISTRIBUTION, HABITAT, AND ASSOCIATED FAUNA (Reed, 2004). Council staff provided an overview of the integration of new deepwater coral HAPCs into the Fishery Ecosystem Plan and Comprehensive Ecosystem Amendment development process. Panel members discussed the information provided to further refine previous recommendations on the establishment of new deepwater coral HAPCs in the South Atlantic Region. In addition, Council staff provided an overview of the preliminary development of a deepwater coral research and monitoring plan
Proposed Deepwater Coral HAPCs The excerpts below are from S. Ross' report and provide a more detailed description of each proposed site off North Carolina.
Cape Lookout Lophelia Bank A: Aside from a few maps there are no published data from this coral mound. Between summer 2000 and summer 2004 Ross et al. (unpubl. data) sampled this area extensively using a variety of methods throughout the water column. Their major method for collecting bottom data on the reef proper was the Johnson Sea Link (JSL) research submersible. Fifteen dives were made on coral mounds in this area and observations from these totaling nearly 33 hours (bottom time) are the basis of the descriptions of habitat and fauna below. Preliminary observations suggest that this area contains the most extensive coral mounds off North Carolina; however, it must be emphasized that data are lacking to adequately judge overall sizes and areal coverage. There appear to be several prominences capping a ridge system, thus, presenting a very rugged and diverse bathymetry, but there are also other mounds away from the main ridge sampled. The main mound system rises vertically nearly 80 m over a distance of about 1 km, and in places exhibits slopes in excess of 50-60 degrees. Sides and tops of these mounds are covered with extensive colonies of living Lophelia pertusa, with few other corals being observed. Dead colonies and coral rubble interspersed with sandy channels are also abundant. Extensive coral rubble zones surround the mounds for a large, but unknown, distance (exact area not yet surveyed), especially at the bases of the mounds/ridges, and in places seem to be quite thick. These topographic highs accelerate bottom currents which favor attached filterfeeders. Because fishes are somewhat disturbed by submersibles, data on the fish community has accumulated slowly; however, this group is quite diverse on the coral habitat. Ross et al. have so far identified over 43 benthic or
- 3 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006 benthopelagic fish species on and around these coral banks. Of the twenty five total fish species occurring on prime coral habitat of Bank A, nine dominate the data. Beryx decadactylus usually occurs in large aggregations moving over the reef, while most other major species occur as single individuals. Many of these species are cryptic, being well hidden deep in the corals (e.g., Hoplostethus occidentalis, Netenchelys exoria, Conger oceanicus). The morid, Laemonema melanurum, is one of the larger fishes abundant at every site with corals. This fish seems to rarely leave the prime reef area. Trash and entangled fishing gear were observed on this reef, suggesting some level of commercial fishing pressure. Initially the most impressive biological aspect of these coral mounds (aside from the corals themselves) was the well developed and abundant invertebrate fauna. We have not yet detected major differences in the invertebrate fauna among the three North Carolina banks; therefore, this paragraph is relevant to all three areas. Galatheid crabs (especially Eumunida picta) and the brisingid basket star (Novodinia antillensis) were particularly obvious, perching high in coral bushes to catch passing animals or filter in the currents. One very different aspect of the North Carolina deep coral habitat compared to the rest of the South Atlantic Bight is the massive numbers of a brittle star (Ophiacantha bidentata) covering both dead and living coral colonies. These are perhaps the most abundant macroinvertebrate on these banks. In places the bottom is covered with huge numbers of several species of anemones. The abundance of filter feeders suggests a food rich habitat.
Cape Lookout Lophelia Bank B: Except for a few maps there are no published data from this coral mound. Between summer 2001 and summer 2004 Ross et al. (unpubl. data) sampled this area using a variety of methods throughout the water column. The Johnson Sea Link (JSL) submersible was the major method for collecting bottom data on the reef proper. Five dives were made on coral mounds in this area, and observations from these totaling 10.4 hours form the basis of the descriptions of habitat and fauna below. The least amount of data are available for this area. Mounds appear to cover a smaller area than those described above, but here again better mapping data are needed. These mounds rise at least 53 m over a distance of about 0.4 km. There is a small mound away from the main system and in general these mounds were less dramatic than those described above. They appeared to be of the same general construction as Bank A, appearing to be built of coral rubble matrix that had trapped sediments. Extensive fields of coral rubble surrounded the area. Both living and dead corals were common on this bank, with some living bushes being quite large. Preliminary analyses (Ross et al. unpubl.) have identified 11 fish species from this bank, but it is clear that the species list would be much higher in this well developed habitat if there were more samples. The dominant fish species appears to be Helicolenus dactylopterus, followed by L. melanurum, H. occidentalis, L. barbatulum, and N. exoria. Although H. dactylopterus can be common on all habitats, it clearly occurs most often around structures. It is intimately associated with the coral substrate, and it is very abundant around this reef habitat. The invertebrate fauna on this reef system does not appear substantially different from Bank A
Cape Fear Lophelia Bank:
- 4 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006
Aside from the map in EEZ-SCAN 87 Scientific Staff (1991) there are no published data from this coral mound and no indication that it was sampled before the studies initiated by Ross et al. (unpubl. data) between summer 2002 and summer 2004. Ross et al. located this bank based on estimated coordinates from the USGS survey (EEZ-SCAN 87 Scientific Staff 1991). As above, the JSL submersible was the major method for collecting bottom data on the reef proper. Seven dives were made on coral mounds in this area, and observations from these totaling 15.4 hours were used to describe the habitat and fauna. Sampling in this area was focused on a relatively small area, but data are lacking to accurately estimate the size and area covered by coral mounds or rubble zones. These mounds rise nearly 80 m over a distance of about 0.4 km, and exhibit some of the most rugged habitat and vertical excursion of any area sampled. This mound system also appears to be of the same general construction as Banks A and B, being built of coral rubble matrix with trapped sediments. Fields of coral rubble are common around the area. Both living and dead corals were common on this bank. The greatest numbers of large fishes were observed on this bank. Twelve total fish species were observed here, but as above, this list should increase with increasing sampling effort. As on Banks A and B, decadactylus was the most common fish, followed closely by Polyprion americanus (wreckfish). So far, of the three North Carolina banks, this is the only area where wreckfish have been observed, and on some dives 8-10 large individuals were seen swimming slowly along the sides of the ridges. However, it is very likely that wreckfish occur on the other banks. As on the other two banks, L. melanurum was common here, always on prime reef habitat. Conger oceanicus (always large adults) and Myxine glutinosa were both frequently observed on this bank. The invertebrate fauna on this reef system does not appear substantially different from Banks A and B.
The following excerpts are from J. Reed's report for proposed HAPC sites off SC, GA and FL.
Region D: Stetson Reefs, Eastern Blake Plateau (from Reed, 2002a; Reed et al., 2004b): This site is on the outer eastern edge of the Blake Plateau, ~120 nm SE of Charleston, South Carolina, at depths of 640-869 m. Over 200 coral mounds up to 146 m in height occur over this 6174 km2 area that was first described by Thomas Stetson from echo soundings and bottom dredges (Stetson et al., 1962; Uchupi, 1968). These were described as steep-sloped structures with active growth on top of the banks. Live coral colonies up to 50 cm in diameter were observed with a camera sled. Enallopsammia profunda (=D. profunda) was the dominant species in all areas although Lophelia pertusa was concentrated on top of the mounds. Densest coral growth occurred along an escarpment at Region D1. Stetson et al. (1962) reported an abundance of hydroids, alcyonaceans, echinoderms, actiniaria, and ophiuroids, but a rarity of large mollusks. The flabelliform gorgonians were also current-oriented. Popenoe and Manheim (2001) have made detailed geological maps of this Charleston Bump region which also indicate numerous coral mounds. Recent fathometer transects by the PI indicated dozens and possibly hundreds of individual pinnacles and mounds within the small region that we surveyed which is only a fraction of the Stetson Bank area (Reed and Pomponi, 2002b; Reed et al., 2002; Reed et al., 2004b). From our fathometer transects, two pinnacle regions were selected. Three submersible dives were made on “Pinnacle 3” and four dives on “Stetson’s Peak” which
- 5 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006 is described below. A small subset of the Stetson Bank area was first mapped during six fathometer transects covering ~28 nm2, in which six major peaks or pinnacles and four major scarps were plotted. The base depth of these pinnacles ranged from 689 m to 643 m, with relief of 46 to 102 m. A subset of this was further mapped with 70 fathometer transects spaced 250 m apart (recording depth, latitude and longitude ~ every 3 seconds), covering an area of 1 x 1.5 nm, resulting in a 3-D bathymetric GIS Arcview map of a major feature, which we named Stetson’s Pinnacle. Stetson’s Pinnacle was 780 m at the south base and the peak was 627 m. This represents one of the tallest Lophelia coral lithoherms known, nearly 153 m in relief. The linear distance from the south base to the peak was ~0.5 nm. The lower flank of the pinnacle from ~762 m to 701 m on the south face was a gentle slope of 10-30o with a series of 3-4 m high ridges and terraces that were generally aligned 60-240o across the slope face. These ridges were covered with nearly 100% Lophelia coral rubble, 15-30 cm colonies of live Lophelia, and standing dead colonies of Lophelia, 30-60 cm tall. Very little rock was exposed, except on the steeper exposed, eroded faces of the ridges. Some rock slabs, ~30 cm thick, have slumped from these faces. From 701 m to 677 m the slope increased from ~45o to 60o. From 671 m to the peak, the geomorphology was very complex and rugged, consisting of 60-90o rock walls and 3-9 m tall rock outcrops. Colonies of Lophelia, 30-60 cm tall, were more common, and some rock ledges had nearly 100% cover of live Lophelia thickets. The top edge of the pinnacle was a 30 cm thick rock crust which was undercut from erosion; below this was a 90o escarpment of 3-6 m. The peak was a flat rock plateau at 625- 628 m and was approximately 0.1 nm across on a S-N submersible transect. The north face was not explored in detail but is a vertical rock wall from the peak to ~654 m then grades to a 45o slope with boulders and rock outcrops. Dominant sessile macrofauna consisted of scleractinia, stylasterine hydrocorals, gorgonacea and sponges. The colonial scleractinia were dominated by colonies of Lophelia pertusa (30-60 cm tall) and Enallopsammia profunda, and Solenosmilia variabilis were present. Small stylasterine corals (15 cm tall) were common and numerous species of solitary cup corals were abundant. Dominant octocorallia consisted of colonies of Primnoidae (15-30 cm tall), paramuriceids (60-90 cm), Isididae bamboo coral (15-60 cm), stolonifera, and stalked Nephtheidae (5-10 cm). Dominant sponges consisted of Pachastrellidae (25 cm fingers and 25- 50 cm plates), Corallistidae (10 cm cups), Hexactinellida glass sponges (30 cm vase), Geodia sp. (15-50 cm spherical), and Leiodermatium sp. (50 cm frilly plates). Although motile fauna were not targeted, some dominant groups were noted. No large decapods crustaceans were common although some red portunids were observed. Two species of echinoids were common, one white urchin and one stylocidaroid. No holothurians or asteroids were noted. Dense populations of Ophiuroidea were visible in close-up video of coral clusters and sponges. No large Mollusca were noted except for some squid. Fish consisted mostly of benthic gadids and rattails. On the steeper upper flank, from 671 to 625 m the density, diversity, and size of sponges increased; 15- 50 cm macro sponges were more abundant. Massive Spongosorites sp. were common, Pachastrellidae tube sponges were abundant, and Hexactinellida glass sponges were also common. On the peak plateau the dominant macrofauna were colonies of Lophelia pertusa (30- 60 cm tall), coral rubble, Phakellia sp. fan sponges (30-50 cm), and numerous other demosponges were abundant. No large fish were seen on top.
- 6 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006
Region C: Savannah Lithoherms, Blake Plateau (from Reed, 2002a; Reed et al., 2004b): A number of high-relief lithoherms occur within this region of the Blake Plateau ~90 nm east of Savannah, Georgia. Region C is at the base of the Florida-Hatteras Slope, near the western edge of the Blake Plateau, and occurs in a region of phosphoritic sand, gravel and rock pavement on the Charleston Bump (Sedberry, 2001). Wenner and Barans (2001) described 15-23 m tall coral mounds in this region that were thinly veneered with fine sediment, dead coral fragments and thickets of Lophelia and Enallopsammia. They found that blackbellied rosefish and wreckfish were frequent associates of this habitat. In general, the high-relief Lophelia mounds occur in this region at depths of 490-550 m and have maximum relief of 61 m. JSL-II dives 1690, 1697 and 1698 reported a coral rubble slope with <5% cover of 30 cm, live coral colonies (Reed, 2002a). On the reef crest were 30-50 cm diameter coral colonies covering ~10% of the bottom. Some areas consisted of a rock pavement with a thin veneer of sand, coral rubble, and 5-25 cm phosphoritic rocks. At Alvin dive sites 200 and 203, Milliman et al. (1967) reported elongate coral mounds, approximately 10 m wide and 1 km long, that were oriented NNE-SSW. The mounds had 25-37o slopes and 54 m relief. Live colonies (10-20 cm diameter) of E. profunda (=D. profunda) dominated and L. pertusa (=L. prolifera) was common. No rock outcrops were observed. These submersible dives found that these lithoherms provided habitat for large populations of massive sponges and gorgonians in addition to the smaller macroinvertebrates which have not been studied in detail. Dominant macrofauna included large plate-shaped sponges (Pachastrella monilifera) and stalked, fan-shaped sponges (Phakellia ventilabrum), up to 90 cm in diameter and height. At certain sites (JSL-II dive 1697), these species were estimated at 1 colony/10 m2. Densities of small stalked spherical sponges (Stylocordyla sp., Hadromerida) were estimated in some areas at 167 colonies/10 m2. Hexactinellid (glass) sponges such as Farrea? sp. were also common. Dominant gorgonacea included Eunicella sp. (Plexauridae) and Plumarella pourtalessi (Primnoidae). Recent fathometer transects by the PI at Savannah Lithoherm Site #1 (JSL II-3327) extended 2.36 nm S-N revealed a massive lithoherm feature that consisted of five major pinnacles with a base depth of 549 m, minimum depth of 465 m, and maximum relief of 83 m (Reed and Pomponi, 2002b; Reed et al., 2002; Reed et al., 2004b). The individual pinnacles ranged from 9 to 61 m in height. A single submersible transect, south to north, on Pinnacle #4 showed a minimum depth of 499 m. The south flank of the pinnacle was a gentle 10-20o slope, with ~90% cover of coarse sand, coral rubble and some 15 cm rock ledges. The peak was a sharp ridge oriented, NW-SE, perpendicular to the prevailing 1 kn current. The north side face of the ridge was a 45o rock escarpment of about 3 m which dropped onto a flatter terrace. From a depth of 499 to 527 m, the north slope formed a series of terraces or shallow depressions, ~9-15 m wide, that were separated by 3 m high escarpments of 30-45\o. Exposed rock surfaces showed a black phosphoritic rock pavement. The dominant sessile macrofauna occurred on the exposed pavement of the terraces and in particular at the edges of the rock outcrops and the crest of the pinnacle. The estimated cover of sponges and gorgonians was 10% on the exposed rock areas. Colonies of Lophelia pertusa (15-30 cm diameter) were common but not abundant with ~1% coverage. Dominant Cnidaria included several species of gorgonacea (15-20 cm tall), Primnoidae, Plexauridae (several spp.), Antipathes sp. (1 m tall), and Lophelia pertusa. Dominant sponges included large Phakellia ventilabrum (fan sponges, 30-90 cm diameter), Pachastrellidae plate sponges (30 cm),
- 7 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006
Choristida plate sponges (30 cm), and Hexactinellid glass sponges. Motile fauna consisted of decapod crustaceans (Chaceon fenneri, 25 cm; and Galatheidae, 15 cm) and mollusks. Few large fish were observed but a 1.5 m swordfish, several 1 m sharks, and numerous blackbelly rosefish were noted. A fathometer transect by the PI at Savannah Lithoherm Site 2 extended 4.6 nm, SW to NE , mapped 8 pinnacles with maximum depth of 549 m and relief of 15-50 m. Submersible dives were made on Pinnacles 1, 5 and 6 of this group. Pinnacle 1 was the largest feature of this group; the base was 537 m and the top was 487 m. The south face, from a depth of 518 to 510 m, was a gentle 10o slope, covered with coarse brown sand and Lophelia coral rubble. A 3-m high ridge of phosphoritic rock, extended NE-SW, cropped out at a depth of 510 m. This was covered with nearly 100% cover of 15 cm thick standing dead Lophelia coral and dense live colonies of Lophelia pertusa (15-40 cm). From depths of 500 m to 495 m were a series of exposed rock ridges and terraces, that were 3-9 m tall with 45o slopes. Some of the terraces were ~30 m wide. Each ridge and terrace had thick layers of standing dead Lophelia, and dense live coral. These had nearly 100% cover of sponges (Phakellia sp., Geodia sp., Pachastrellidae, and Hexactinellida), scleractinia (Lophelia pertusa, Madrepora oculata), stylasterine hydrocorals, numerous species of gorgonacea (Ifalukellidae, Isididae, Primnoidae), and 1 m bushes of black coral (Antipathes sp.). Deep deposits of sand and coral rubble occurred in the depressions between the ridges. The north face, from 500 m to 524 m was a gentle slope of 10o, that had deep deposits of coarse brown foraminiferal sand and coral rubble. Exposed rock pavement was sparse on the north slope, but a few low rises with live bottom habitat occurred at 524 m. Dominant mobile fauna included decapod crustaceans (Chaceon fenneri, 15 cm Galatheidae), rattail fish, and 60 cm sharks were common.
Region B: Florida Lophelia Pinnacles (from Reed, 2002a; Reed et al., 2004b) Numerous high-relief Lophelia reefs and lithoherms occur in this region at the base of the Florida- Hatteras Slope and at depths of 670-866 m. The reefs in the southern portion of this region form along the western edge of the Straits of Florida and are 15-25 nm east of the Oculina coral banks Marine Protected Area (MPA). Along a 222-km stretch off northeastern and central Florida (from Jacksonville to Jupiter), nearly 300 mounds from 8 to 168 m in height (25- 550 ft) were recently mapped by the PI using a single beam echosounder (Fig. 11; Reed et al., 2004b). Between 1982 and 2004, dives with the Johnson-Sea-Link (JSL) submersibles and ROVs by the PI confirmed the presence of Lophelia mounds and lithoherms in this region (Reed, 2002a; Reed et al., 2002; Reed and Wright, 2004; Reed et al., 2004b). The northern sites off Jacksonville and southern Georgia appeared to be primarily lithoherms which are pinnacles capped with exposed rock (described in part by Paull et al., 2000), whereas the features from south of St. Augustine to Jupiter were predominately Lophelia coral pinnacles or mud mounds capped with dense 1-m-tall thickets of Lophelia pertusa and Enallopsammia profunda with varying amounts of coral debris and live coral. Dominant habitat-forming coral species were Lophelia pertusa, Madrepora oculata, Enallopsammia profunda, bamboo coral (Isididae), black coral (Antipatharia), and diverse populations of octocorals and sponges (Reed et al., 2004b). Paull et al. (2000) estimated that over 40,000 coral lithoherms may be present in this region of the Straits of Florida and the Blake Plateau. Their dives with the Johnson-Sea-Link submersible and the U.S. Navy’s submarine NR-1
- 8 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006 described a region off northern Florida and southern Georgia of dense lithoherms forming pinnacles 5 to 150 m in height with 30-60o slopes that had thickets of live ahermatypic coral (unidentified species, but photos suggest Lophelia and/or Enallopsammia). The depths range from 440 to >900 m but most mounds were within 500-750 m. Each lithoherm was ~100-1000 m long and the ridge crest was generally oriented perpendicular to the northerly flowing Gulf Stream current (25-50 cm s-1 on flat bottom, 50-100 cm s-1 on southern slopes and crests). Thickets of live coral up to 1 m were mostly found on the southern facing slopes and crests whereas the northern slopes were mostly dead coral rubble. These were termed lithoherms since the mounds were partially consolidated by a carbonate crust, 20-30 cm thick, consisting of micritic wackestone with embedded planktonic foraminifera, pteropods, and coral debris (Paull et al., 2000). A recent echosounder transect by the PI revealed a massive lithoherm, 3.08 nm long (N-S) that consisted of at least 7 individual peaks with heights of 30-60 m (Fig. 12; Reed and Wright, 2004; Reed et al., 2004b). The maximum depth was 701 m with total relief of 157 m. Three submersible dives (JSL II-3333, 3334; I-4658) were made on Peak 6 of pinnacle #204B which was the tallest individual feature of the lithoherm with maximum relief of 107 m and a minimum depth at the peak of 544 m (Reed et al., 2004b). The east face was a 20-30o slope and steeper (50o) near the top. The west face was a 25-30o slope which steepened to 80o from 561 m to the top ridge. The slopes consisted of sand and mud, rock pavement and rubble. A transect up the south slope reported a 30-40o slope with a series of terraces and dense thickets of 30-60 cm tall dead and live Lophelia coral that were mostly found on top of mounds, ridges and terrace edges. One peak at 565 m had dense thickets of live and dead standing Lophelia coral (~20% live) and outcrops of thick coral rubble. Dominant sessile fauna consisted of Lophelia pertusa, abundant Isididae bamboo coral (30-60 cm) on the lower flanks of the mound, Antipatharia black coral, and abundant small octocorals including the gorgonacea (Placogorgia sp., Chrysogorgia sp, and Plexauridae) and Nephtheidae soft corals (Anthomastus sp., Nephthya sp.). Dominant sponges consisted of Geodia sp., Phakellia sp., Spongosorites sp. Petrosiidae, Pachastrellidae, and Hexactinellida. Further south off Cape Canaveral, echosounder transects by the PI on Lophelia Pinnacle #113 revealed a 61 m tall pinnacle with maximum depth of 777 m. The width (NW-SE) was 0.9 nm and consisted of at least 3 individual peaks or ridges on top, each with 15-19 m relief. One submersible dive (JSL II-3335) reported 30-60o slopes, with sand, coral rubble, and up to 10% cover of live coral. No exposed rock was observed. This appeared to be a classic Lophelia mud mound. The second dive site (JSL II-3336) at Pinnacle #151 was also a deep-water Lophelia coral reef comprised entirely of coral and sediment. Maximum depth was 758 m, with 44 m relief, and ~0.3 nm wide (N-S). The top was a series of ridged peaks from 713 to 722 m in depth. The lower flanks of the south face was a 10-20o slope of fine light colored sand with a series of 1-3 m high sand dunes or ridges that were linear NW-SE. The ridges had ~50% cover of thickets of Lophelia pertusa coral. The thickets consisted of 1 m tall dead, standing and intact, Lophelia pertusa colonies. Approximately 1-10% were alive on the outer parts (15-30 cm) on top of the standing dead bases. There was very little broken dead coral rubble in the sand and there was no evidence of trawl or mechanical damage. Most of the coral was intact, and the dead coral was brown. The sand between the ridges was fine and light colored, with 7-15 cm sand waves. The upper slope steepened to 45o and 70-80o slope near the upper 10 m from the top. The top of the
- 9 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006 pinnacle had up to 100% cover of 1-1.5 m tall coral thickets, on a narrow ridge that was 5-10 m wide. The coral consisted of both Lophelia pertusa and Enallopsammia profunda. Approximately 10-20% cover was live coral of 30-90 cm. The north slope was nearly vertical (70-80o) for the upper 10 m then consisted of a series of coral thickets on terraces or ridges. No exposed rock was visible and the entire pinnacle appeared to be a classic Lophelia mud mound. No discernable zonation of macrobenthic fauna was apparent from the base to the top. Corals consisted of Lophelia pertusa, Enallopsammia profunda, Madrepora oculata, and some stylasterine hydrocorals. Dominant octocoral gorgonacea included Primnoidae (2 spp.), Isididae bamboo coral (Isidella sp. and Keratoisis flexibilis), and the alcyonaceans Anthomastus sp. and Nephthya sp. Dominant sponges consisted of several species of Hexactinellida glass sponges, large yellow demosponges (60-90 cm diameter), Pachastrellidae, and Phakellia sp. fan sponges. Echinoderms included urchins (cidaroid and Hydrosoma? sp.) and comatulid crinoids, but no stalked crinoids. Some large decapod crustaceans included Chaceon fenneri and large galatheids. No mollusks were observed but were likely within the coral habitat that was not collected. Common fish were 2 m sharks, 25 cm eels, 25 cm skates, chimaera, and blackbelly rosefish.
Region G: The Miami Terrace Escarpment (from Reed et al., 2004b) The Miami Terrace is a 65-km long carbonate platform that lies between Boca Raton and South Miami at depths of 200-400 m in the northern Straits of Florida. It consists of high- relief Tertiary limestone ridges, scarps and slabs that provide extensive hard bottom habitat (Uchupi, 1966, 1969; Kofoed and Malloy, 1965; Uchupi and Emery, 1967; Malloy and Hurley, 1970; Ballard and Uchupi, 1971; Neumann and Ball, 1970). At the eastern edge of the Terrace, a high-relief, phosphoritic limestone escarpment of Miocene age with relief of up to 90 m at depths of 365 m is capped with Lophelia pertusa coral, stylasterine hydrocoral (Stylasteridae), bamboo coral (Isididae), and various sponges and octocorals (Reed et al., 2004b; Reed and Wright, 2004). Dense aggregations of 50-100 wreckfish were observed here by the PI during JSL submersible dives in May 2004 (Reed et al., 2004b). Previous studies in this region include geological studies on the Miami Terrace (Neumann and Ball, 1970; Ballard and Uchupi, 1971) and dredge- and trawlbased faunal surveys in the 1970s primarily by the University of Miami (e.g., Halpern, 1970; Holthuis, 1971, 1974; Cairns, 1979). Lophelia mounds are also present at the base of the escarpment (~670 m) within the axis of the Straits of Florida, but little is known of their distribution, abundance or associated fauna. Using the Aluminaut submersible, Neumann and Ball (1970) found thickets of Lophelia, Enallopsammia (=Dendrophyllia), and Madepora growing on elongate depressions, sand ridges and mounds. Large quantities of L. pertusa and E. profunda have also been dredged from 738-761 m (Cairns, 1979). Recent JSL submersible dives and fathometer transects by the PI at four sites (Reed Site #BU4, 6, 2, and 1b) indicated the outer rim of the Miami Terrace to consist of a double ridge with steep rocky escarpments (Table 1; Fig. 6; Reed and Wright, 2004; Reed et al., 2004b). At Miami Terrace Site #BU4, the narrow N-S trending east ridge was 279 m at the top and had a steep 95 m. escarpment on the west face. The east and west faces of the ridges were 30-40o slopes with some near vertical sections consisting of dark brown phosphoritic rock pavement, boulders and outcrops. The crest of the east ridge was a narrow plateau ~10 m wide. At Site #BU6, the crest of
- 10 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006 the west ridge was 310 m and the base of the valley between the west and east ridges was 420 m. At Site #BU2, the echosounder transect showed a 13 m tall rounded mound at a depth of 636 m near the base of the terrace within the axis of the Straits of Florida. The profile indicated that it is likely a Lophelia mound. West of this feature the east face of the east ridge was a steep escarpment from 567 m to 412 m at the crest. The west ridge crested at 321 m. Total distance from the deep mound to the west ridge was 2.9 nm. Site #BU1b was the most southerly transect on the Miami Terrace. An E-W echosounder profile at this site indicated a double peaked east ridge cresting at 521 m, then a valley at 549 m, and the west ridge at 322 m. The east face of the west ridge consisted of a 155 m tall escarpment. There were considerable differences among the sites in habitat and fauna; however, in general, the lower slopes of the ridges and the flat pavement on top of the terrace were relatively barren. However, the steep escarpments especially near the top of the ridges were rich in corals, octocorals, and sponges. Dominant sessile fauna consisted of the following Cnidaria: small (15- 30 cm) and large (60-90 cm) tall octocoral gorgonacea (Paramuricea spp., Placogorgia spp., Isididae bamboo coral); colonial scleractinia included scattered thickets of 30-60 cm tall Lophelia pertusa (varying from nearly 100% live to 100% dead), Madrepora oculata (40 cm), and Enallopsammia profunda; stylasterine hydrocorals (15-25 cm); and Antipatharia (30-60 cm tall). Diverse sponge populations of Hexactinellida and Demospongiae included: Heterotella sp., Spongosorites sp., Geodia sp., Vetulina sp., Leiodermatium sp., Petrosia sp., Raspailiidae, Choristida, Pachastrellidae, and Corallistidae. Other motile invertebrates included Asteroporpa sp. ophiuroids, Stylocidaris sp. urchins, Mollusca, Actiniaria, and Decapoda crustaceans (Chaceon fenneri and Galatheidae). Schools of ~50-100 wreckfish (Polyprion americanus), ~60-90 cm in length, were observed on several submersible dives along with blackbelly rosefish, skates, sharks, and dense schools of jacks.
Region H: Portales Terrace Lithoherms (from Reed et al., 2004a) The Pourtalès Terrace provides extensive, high-relief, hard-bottom habitat, covering 3,429 km2 (1,000 nm2) at depths of 200-450 m. The Terrace parallels the Florida Keys for 213 km and has a maximum width of 32 km (Jordan, 1954; Jordan and Stewart, 1961; Jordan et al., 1964; Gomberg, 1976; Land and Paull, 2000). Reed et al. (2004a) surveyed several deep-water, high-relief, hardbottom sites including the Jordan and Marathon deep-water sinkholes on the outer edge of the Terrace, and five high-relief bioherms on its central eastern portion. The JSL and Clelia submersibles were used to characterize coral habitat and describe the fish and associated macrobenthic communities. These submersible dives were the first to enter and explore any of these features. The upper sinkhole rims range from 175 to 461 m in depth and have a maximum relief of 180 m. The Jordan Sinkhole may be one of the deepest and largest sinkholes known. The high- relief area of the middle and eastern portion of the Pourtalès Terrace is a 55 km-long, northeasterly trending band of what appears to be karst topography that consists of depressions flanked by well defined knolls and ridges with maximum elevation of 91 m above the terrace (Jordan et al., 1964; Land and Paull, 2000). Further to the northeast of this knoll-depression zone is another zone of 40-m high topographic relief that lacks any regular pattern (Gomberg, 1976). The high-relief bioherms (the proposed HAPC sites within this region) lie in 198 to 319 m, with a maximum height of 120 m. A total of 26 fish taxa were identified from the sinkhole and bioherm sites (Table 4). Species of
- 11 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006 potential commercial importance included tilefish, sharks, speckled hind, yellow-edge grouper, warsaw grouper, snowy grouper, blackbelly rosefish, red porgy, drum, scorpion fish, amberjack, and phycid hakes. Many different species of Cnidaria were recorded, including Antipatharia black corals, stylasterine hydrocorals, octocorals, and one colonial scleractinian (Solenosmilia variabilis).
Tennessee and Alligator Humps, Bioherms #1-4- Pourtalès Terrace (from Reed et al., 2004a) The Tennessee and Alligator Humps are among dozens of lithoherms that lie in a region called “The Humps” by local fishers, ~14 nm south of the Florida Keys and south of Tennessee and Alligator Reefs. Three dives were made by the PI on Bioherm #3 (Clelia 597, 598, 600; Aug. 2001), approximately 8.5 nm NE of Bioherm#2 (Fig. 15). Bioherm #3 consisted of two peaks 1.05 nm apart with a maximum relief of 62 m. The North Peak’s minimum depth was 155 m and was 653 m wide at the base, which was 217 m deep at the east base and 183 m at the west side. The minimum depth of South Peak was 160 m and was about 678 m in width E to W at the base. The surrounding habitat adjacent to the mounds was flat sand with about 10% cover of rock pavement. From 213 m to the top, generally on the east flank of the mound, were a series of flat rock pavement terraces at depths of 210, 203, 198, 194, 183, and 171 m and the top plateau was at 165 m. Between each terrace a 30-45o slope consisted of either rock pavement or coarse sand and rubble. Below each terrace was a vertical scarp of 1-2 m where the sediment was eroded away leaving the edge of the terrace exposed as a horizontal, thin rock crust overhang of <1 m and 15-30 cm thick. The top of the bioherm was a broad plateau of rock pavement with 50-100% exposed rock, few ledges or outcrops, and coarse brown sand. Less time was spent on the western side, which was more exposed to the strong bottom currents. The west side of South Peak sloped more gradually than the eastern side, had more sediment, and no ledges were observed.
Fish Communities (from Reed et al., 2004a) A total of 31 fish taxa, of which 24 were identified to species level, were identified from our submersible videotapes and were associated with the deep-water sinkholes and high- relief bioherms. Few studies have directly documented deep-water fish associations with deep-water reef habitats in the western Atlantic. Most of the work has concentrated on the Charleston Bump region of the Blake Plateau off Georgia and South Carolina (Sedberry, 2001). Ross (pers. comm.) reported the following species are common to both the deep-water Lophelia reefs on the Blake Plateau off the Carolinas and those of this study: Chloropthalmus agassizi, Helicolenus dactylopterus, Hoplostethus sp., Laemonema melanurum, Nezumia sp., and Xiphias gladius. Species most common to the high-relief bioherms included deepbody boarfish, blueline tilefish, snowy grouper, and roughtongue bass. Some species were common at both the sinkhole and bioherm sites and included snowy grouper, blackbelly rosefish, and mora. In addition to the moribund swordfish observed in the Jordan Sinkhole, a swordfish was observed from the NR-1 submersible on top of Pourtales Terrace (C. Paull, pers. observation). Species of potential commercial importance included tilefish, sharks, speckled hind, yellowedge grouper, warsaw grouper, snowy grouper, blackbelly rosefish, red porgy, drum, scorpionfish, amberjack, and phycid hakes. However, the fish densities that we saw at any of the sites were in insufficient numbers to suggest commercial or recreation harvest. In fact, any of
- 12 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006 the features, both sinkholes and bioherms, could be overfished very easily since only a few individuals of the larger grouper species were present at any one site.
Benthic Communities (from Reed et al., 2004a) The benthos at the bioherm sites was dominated by sponges, octocorals and stylasterids. A total of 21 taxa of Cnidaria were sampled or observed and 16 were identified to species level. These included 3 species of antipatharian black coral, 5 stylasterid hydrocorals, 11 octocorals with one possible new species, and 1 scleractinian (Solenosmilia variabilis). Eight species were associated only with the Pourtalès sinkholes and not the bioherms; these included two species of antipatharians; the octocorals Paramuricea placomus, Plumarella pourtalesii, Trachimuricea hirta; and the scleractinian Solenosmilia variabilis. Although Gomberg (1976) found evidence of skeletal remains of the colonial scleractinians Lophelia and Madrepora in sediment samples from the terrace, we did not see any colonies at our dive sites. Sponges identified from collections included 28 taxa. Five species of stylasterine hydrocorals were Distichopora foliacea, Pliobothrus echinatus, Stylaster erubescens, S. filogranus, and S. miniatus. On the flat pavement adjacent to the base of the mounds, stylasterids and antipatharian black coral bushes were common along with sea urchins and sea stars. The densities of sponges, stylasterid hydrocorals and octocorals were very high, especially on the plateaus and terraces of the bioherms on the Pourtalès Terrace. Maximum densities of sponges (>5 cm) on the plateaus ranged from 1-80 colonies m-2. Stylasterid coral densities ranged from 9-96 colonies m-2 and octocorals 16-48. Densities of sponges (1-2 colonies m-2) and stylasterids (1-20) also dominated the terraces and slopes of the bioherm sites but generally in lower densities than the peak plateaus whereas the octocorals generally had higher densities on the flanks (1-80 colonies m-2).
2006 Updates to the Council and Advisory Panels Updated reports on deepwater coral habitat distribution and characterization in the South Atlantic Region were presented by John Reed with the Harbor Branch Oceanographic Institute and Steve Ross with UNCW/USGS. That updated information was used to formulate modifications to the proposed deepwater Coral HAPCs.
JOINT HABITAT AND CORAL AP RECOMMENDATIONS
Proposed Deepwater Coral HAPCs: • The above proposed deepwater coral HAPCs should be expanded based on new research and data compiled for the Council and presented to the Advisory Panels by John Reed and Steve Ross. Specifically, the large central area should be expanded to connect Stetson Reefs, enlarge somewhat to the north to include newly documented sites and enlarged west to include the 400 meter isobath.
• The large central area should be connected with the Miami Terrace C-HAPC, also using the 400 meter isobath as the western boundary.
• The Miami Terrace C-HAPC should be expanded to the edge of the EEZ to the east (to include mound and pinnacle structures that extend toward and into the
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Bahamian EEZ. The western boundary should be extended to include the 300 meter isobath to include newly documented deep coral habitat.
• Expand the Portales Terrace C-HAPC to cover newly documented deepwater coral habitat.
• Recognizing that deepwater ecosystems are not closed, and do have connections internationally, the Panels request the Council interact with the Bahamian government and Department of State, to work with them to find ways to collaborate on research as well as protection measures. The Council could communicate with the Bahamian government directly or through the U.S. Departments of Commerce and Department of State.
Regulations in proposed deepwater Coral HAPCs: The original recommendations by the Advisory Panels are restated as follows: Recommended management measures in all the deepwater coral HAPC sites include the following: prohibit all bottom-disturbing activities, prohibit harvest of corals, and compile a list of threats. The intent would be to prevent any allowable harvest presently permitted under the coral plan, in any deepwater coral HAPC. To prohibit the collection of gorgonians in coral HAPCs - clarify the prohibition would not apply to biomedical or taxonomic collections. To prohibit any type of anchoring. To identify the potential damage associated with other bottom gears (e.g., a future research priority - if damage occurs with the use of planers and cannonball weights).
The Panels reaffirmed their recommendation that damaging gear be precluded. In addition, the Panels requests the Council consider establishing allowable gear to identify appropriate, non-damaging gears. Non-fishing impacts would be fully covered in the Fishery Ecosystem Plan and in future habitat policy statements.
Development of a Deepwater Coral Research and Monitoring Plan for the South Atlantic Region The Habitat and Coral Advisory Panels fully endorse the completion and full implementation of a Deepwater Coral Research and Monitoring Plan.
Development of Rapid Assessment Tool (SEADESC) and Integration into Habitat and Ecosystem IMS The Habitat and Coral Advisory Panels strongly endorses the completion of the processing of existing SEADESC information and presentation in the IMS.
3) Sargassum Research and Management The Habitat and Coral Advisory Panels after being presented a summary of research conducted since the implementation of the Sargassum FMP made the following recommendations based on the complexity of this pelagic habitat and its role as EFH.
JOINT HABITAT AND CORAL AP RECOMMENDATIONS Establish zero harvest of Sargassum through the Comprehensive Ecosystem Amendment.
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Researchers would be allowed to sample under scientific collecting permits with appropriate reporting.
4) Listing of Elkhorn and Staghorn Corals as Threatened under ESA After receiving a briefing on NOAA’s recent decision to go forward with listing of these two species as threatened under ESA the Habitat and Coral Advisory Panels made the following recommendations.
JOINT HABITAT AND CORAL AP RECOMMENDATIONS • The Habitat and Coral Advisory Panels work with NOAA Fisheries, during the 4(d) rulemaking, to identify appropriate conservation measures for inclusion in the recovery plan.
• Work to address ecological problems and threats (including the recovery of Diadema) and maintain genetic diversity is highest. Some of the fledgling restoration operations that are underway need more funding.
• Education needs to be a huge component of this effort with a simple message, such as “Don’t mess with the coral,” being very effective. Funding for education, is just not enough.
• Letters from the Council can make a difference with those receiving them, so the Council should work with the AP to prepare comments as appropriate during the NOAA rulemaking process for Acropora listing and that any comments or recommendations be fully captured in the FEP.
5) Fishery Ecosystem Plan and Comprehensive Ecosystem Amendment After receiving a briefing on the status of the FEP and CEA the Habitat and Coral Advisory Panels made the following recommendations.
JOINT HABITAT AND CORAL AP RECOMMENDATIONS The Advisory Panels strongly endorse continuing movement toward ecosystem based management through the development of the FEP.
The Fishery Ecosystem Plan should: • Cover the tremendous transition that is taking place as small fishing villages are being destroyed by development. The Council has charged the Social Science subcommittee with developing the data that we need to address the threats and challenges to working waterfronts and a workshop to address this is coming up;
• Include a good economic evaluation;
• Quantify ecosystem services;
• Provide a link to each existing ESA Recovery Plan;
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• Accurately characterize fisheries (e.g., Atlantic Menhaden purse seine fishery no longer exists in North Carolina.);
• Provide information on how offshore shoals provide EFH. Applications for alteration of offshore shoals have been submitted and a workshop on how they provide EFH is scheduled in several weeks. Such areas off North Carolina are important during the wintertime for striped bass, Atlantic sturgeon and other species. In addition, offshore soft substrates are import habitat for polychaetes and other species. Federal agency partners are working on the passage through FERC- licensed facilities, but there are many others that aren’t federally-licensed, and use the state priority lists for dam removal, where they exist, in the FEP to recommend some priorities.
• Update, revise and include the Council’s water flow policy, written in 2004, especially to address the Roanoke and Savannah Rivers.
JOINT HABITAT AND CORAL AP CONCENSUS RECOMMENDATIONS: Panel members were requested to provide comments on the potential list of actions for consideration in a developing Comprehensive Ecosystem Amendment. Discussions revolved around the existing list of proposed measures for the Amendment and the following recommended modifications are:
Comprehensive Amendment measures should: • Establish and protect expanded deepwater Coral HAPCs;
• Establish a zero harvest for Sargassum;
• Address octocorals harvest and quota level while considering octocorals as EFH;
• Establish provisions to allow for the discovery of new octocorals species and new compounds (biomedical products), but not for mass exploitation and harvesting of species; and
• Consider invasive species highlighting lionfish in FEP and proposed or future Ecosystem Amendment.
6) Snapper Grouper Amendment 14 - Deepwater Snapper Grouper Marine Protected Areas The Habitat and Coral Advisory Panels were presented with the current proposed alternatives for the establishment of Marine Protected Areas for deepwater snapper grouper species and made the following recommendations.
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Recommendations of the Coral and Habitat and Environmental Protection APs:
1. The sites endorsed originally by these APs captured more valuable habitat than the alternatives added later.
2. Moving sites off-shelf greatly reduces fishery and habitat conservation value by failing to protect juvenile habitat and that for other snapper-grouper species using the shelf break, where place-based management is especially invaluable. Failure to include shelf-break reefs in the deepwater MPAs will necessitate future actions in nearby locations.
3. Sites ultimately approved should contain significant quantities of high value habitat. Many of the sites reviewed did not have adequate survey data to fully characterize them.
4. Sites ultimately approved should be managed to ensure: a) effective education, b) effective outreach, c) effective enforcement, and d) adequate research and monitoring.
5. Adequate resources must be found for this task. Lack of adequate resources will doom this initiative.
6. Implementation of these recommendations must consider the impacts on displaced fishermen and coastal communities.
7. Site-Specific Recommendations:
Snowy Wreck Deepwater MPA: Alternative 1 is preferred because it contains more of the target habitat. Both alternatives contain relatively little of the target habitat. If the focus is the Snowy Wreck itself, a separate much smaller box can be drawn around the wreck. If other alternatives are proposed they should capture more high-value reef habitat.
Northern South Carolina MPA: Alternative 2 is preferred to Alternative 1 because it contains more of the target habitat and better data on reef fish reproduction. Alternative 3 has significantly less value than either Alternative 2 or 1. The eastern half of Alternative 2 has little to no data.
Central South Carolina MPA (Edisto MPA): Alternative 1 is highest priority based on available information, including recent 2005 survey data. Alternative 2 has much less habitat.
Charleston Deep Reef MPA: The alternative is not supported because there is no evidence of appropriate high-value habitat in the site. If the sole purpose is to establish a deepwater artificial reef, the site should be surveyed prior to placement of any material, to verify that existing high value habitat in that site will not be damaged.
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Georgia MPA (Tilefish MPA): Alternative 1 is preferred because it provides significant tilefish habitat and evidence of snowy grouper use. Neither alternative has abundant, documented reef habitat.
North Florida MPA (Mayport MPA/St. Augustine): Alternative 1 is strongly preferred based on the available fish and habitat data. Alternatives 2 and 4 have much less documented high value habitat. The others are unacceptable based on the lack of evidence for high-value habitat and fish use.
Sea Bass Rocks MPA (St. Lucie Hump): This alternative is strongly endorsed based on evidence of high-value habitat, including Oculina coral, and fish usage.
Florida East Hump MPA: This site is strongly endorsed based on data indicating high-value habitat and use by target species. It also includes deepwater coral habitats recommended by the APs for C- HAPC designation.
Vessel Monitoring Systems The Advisory Panels endorsed strong and effective enforcement but felt that other APs (e.g., Enforcement) were in a better position to recommend specific measures.
7) Updated SAFMC Energy Policy Statement Myra Brouwer with Council staff made a presentation highlighting the revisions to the Energy Policy completed through a coordinated effort including Council staff, Jocelyn Karazsia NOAA Fisheries Habitat Conservation Division, Maggie Sloan an intern with Environmental Defense and the Habitat and Coral Advisory Panels.
The Advisory Panels endorse enhancement of the policy to address wind and wave energy facilities, nuclear power cooling water and burgeoning LNG facilities. The Panels also expressed concern about increasing pressures to privatize public trust resources (including the ocean bottom) and conflicts likely to result including conflicts with mandates other than those established under Federal fisheries law.
8) Liquified Natural Gas (LNG) Pipeline Development: Assessing Impacts on Nearshore and Deepwater Coral Habitats Jocelyn Karaszia with NOAA Fisheries Habitat Conservation gave an overview of LNG facilities. NOAA Fisheries is reviewing three LNG facilities. The southernmost project has been authorized by the Corps, but all three projects are awaiting approval from the Bahamian government. Jocelyn noted there are three lines of reefs off south Florida what would have to be crossed to enter at Port Everglades, FL. Jocelyn noted that some of the pipeline would cross an area of previously disturbed habitat, which does support some reef. The applicants propose to use directional drilling to go under reef habitat. Jocelyn reviewed the NMFS concerns, such as punch outs, and release of bentonite drilling muds. Frac-outs are a concern, when the drilling head moves through unconsolidated sediments. Frac-outs can occur anywhere along a route. Wilson asked
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Jocelyn to clarify what a frac-out was. She did so. It is when drilling muds are released to the surface through fissures in the rock, or through unconsolidated sediments. Horizontal Directional Drilling is a 24-hour a day operation. Jocelyn noted that tunneling was proposed, instead of HDD, in order to avoid frac-out potential. Jocelyn noted that tunneling does increase costs, but it is competitive when compared to the total costs including monitoring required for HDD. Tunneling greatly reduced potential impacts. Jocelyn noted the applicant has filed an application with the USCG, for an offshore port for tankers, since the Bahamian government hasn’t approved sites based in that country. Jocelyn reviewed the infrastructure that would be associated with an offshore facility. There would be a thermal plume associated with water used for engine cooling. Larval impingement and entrainment are being investigated. The Calypso applicant plans to begin larval monitoring in July. NMFS has recommended five years of pre-project ichthyoplankton monitoring. Calypso proposes to use a water-glycol mix for warming the gas. Jocelyn noted that one of the offshore terminals is proposed to be sited in one of our recommended HAPCs. She noted that Calypso did assemble an outstanding crew (John Reed, Sandra Brooke among them) to characterize the habitats. Jocelyn noted that a shipping route runs through the middle of the proposed site, which is also highly used by swordfishermen and other users. Jocelyn noted that MARAD is the lead agency for licensing deepwater ports. The Maritime Act of 2002 added LNG to the MARAD mandate. Jocelyn reviewed the permitting process for these facilities, which has a statutory time limit of 365 days. Jocelyn reviewed the NOAA LNG documents and noted they are on the Panel members CD. She reviewed the next steps. NOAA-HCD is coordinating with NURC in Wilmington and with NMFS-SEFSC. Calypso proposes to re-submit its application in July for a USCG “completeness determination.” The completeness determination triggers the NEPA review and 330-day time clock.
9) Research Associated with Proposed LNG Pipeline Development John Reed noted that he was involved in three of the pipeline route proposals. He noted that the reports are not ready yet for release, but he wanted to give us background on what he did for the surveys. He made it clear in his contract, when he was contacted by the consulting firm, for the Seafarer route from the Bahamas to Florida with the Johnson- Sea Link used to survey the route. Where they found hard bottom, it was coral. John Reed told the consultant that his final reported, unaltered would have to go to Florida DEP, and NOAA and the Council. Once it is distributed by Seafarer to the Council, John will discuss it with us. John indicated that the protocol given to them by Florida DEP, NOAA Fisheries and MMS. They used a submersible to fly along the bottom and take complete photo and video documentation. There was a pair of lasers pointing down to allow quantification of the organisms. Percent cover (hard versus soft) bottom, as well as all organisms over three inches were counted. They also have detailed CTD and navigation data for each dive. With respect to the Seafarer pipeline, they did find hard bottom in a short portion of the 18 mile route. A three-mile area had hard-bottom habitat. John mentioned one site, in the first three miles, that had Lophelia, with anywhere from 3-15 foot mounds of coral. There was very little live coral, it was mostly standing dead. John quantified the heights of ridges and so forth. After the first three miles it was just plain mud. There was a lot of trash probably from cruise ships. The Calypso Port Project survey covered 24 square miles of bottom with an ROV. The proposed port would be on
- 19 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006 top of the Miami Terrace, pretty much outside the proposed HAPC we discussed yesterday, but the new deepwater HAPC we discussed would be impacted. John noted that the applicant tried to move their proposed mooring buoys to the west to avoid hard bottom. John noted where there is hard bottom it is low density with few organisms. He noted that two other proposed facilities will cross the entire deepwater HAPC that we are proposing. That area is entirely hard bottom for both the Cliffs of Suez and the AEC proposals. Once the terrace is crossed, you do hit a mud zone, then enter a Lophelia zone, which is close to the EEZ. John’s sites 22, 23 and 59 are the areas where live bottom occurs, Lophelia. There would be varying degrees of impact from the 36-inch pipe. Panel members raised the issue of whether lobster migrations be affected.
The Panels expressed strong support for agency and academic work to protect EFH from damage induced by LNG pipelines and related facilities.
10) Status Report- Consideration of Windfarm Development in the South Atlantic Region Roger Pugliese presented a brief summary of available information on wind development in the South Atlantic and noted to date that there is no formal permit proposal to date. During discussions, they were trying to identify areas offshore, beyond 15 miles. Roger noted there are habitat implications. He noted that if there is a mandate to put all coastal communities at 20 percent of their energy derived from wind, that is a significant potential increase and we should be thinking now about the long-term impacts. Roger reviewed potential fishery impacts. Roger noted there are a lot of data gaps regarding impacts on benthic and pelagic fish fauna. Roger noted there is work being done in NC to evaluate wind potential. He noted the biggest thing is to keep this on the table for consideration in ongoing policy development discussions.
11) Initiation of the Development of a SAFMC Offshore Aquaculture Policy The Advisory Panels received written briefing materials and comment letters from interested parties.
The Panels continue to feel that adoption of this policy is especially important given the agency impetus behind marine aquaculture and intends to complete the drafting of the proposed offshore aquaculture policy before September. The Panels will do so using the Council’s interactive web portal.
12) Invasive Species: Lionfish Research and Proposed Workshop Liz Fairey noted that she was making this presentation on behalf of the NOAA Aquatic Nuisance Species program, although she is housed in NOAA-Fisheries. She noted that the work she would be discussing has been conducted largely by Paula Whitfield, James Morris and Wilson Freshwater.
Two different species have been documented off the coast, but volitans is the major one. The lionfish have been spreading relatively rapidly and the area of potential habitat is
- 20 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006 very large. At 11 C, most of the lionfish die. At 16 C, they stop feeding in the lab, but they haven’t observed cessation of feeding in the wild. Liz noted the researchers are unable to say much about whether the population is increasing, or not, but hope to do so this year. Lionfish have been found in 150-270 feet, which pretty surprising. The sites surveyed included most of the MPA sites. Liz noted this information was presented at an International Aquatic Nuisance Species conference earlier this year. Slides presented showed this species release which was a novelty, has expanded to being the second or third most prevalent fish in formal surveys. Genetics work by Dr. Freshwater indicates the Atlantic Ocean fish have three different haplotypes, which indicates three founder females. The lionfish lay floating eggs, in balls, about 30,000 eggs per spawn. Females mature at around 200-220 mm. For males, maturity is much earlier. They reach maturity at age 1-2. They are eating primarily fishes, but also stomatopod and decapod crustaceans, bivalve and cephalopods and brittle stars. The major prey consumed by the fish analyzed was serranids, followed by scaridae. Doug asked if Liz knew what species of serranids. Stable isotopic ratio analysis suggests that the lionfish are broad generalist feeders. There is a great deal of potential dietary overlap with many native grouper species. Potential threats include human health risk; negative impacts on reef fishes through prey and habitat competition; combined effects to ecosystem from climate change and overfishing; and Caribbean and Gulf expansion potential. The project was funding jointly by NURC and NOAA and NOAA would like to collaborate with the Council on have a lionfish workshop possibly in July, bringing together key researchers and managers to discuss how to deal with the lionfish invasion on the east coast. Liz stated their distribution is really regulated by temperature, and when you get to SC and NC, it is really driven by that factor. Ones driven further north don’t survive. Liz speculated that ones moving inshore would not survive the cold winter temperatures.
The Advisory Panels strongly supported the Council’s engagement through the FEP in addressing ecological implications of invasive species including lionfish, including co- sponsoring the upcoming lionfish workshop.
13) Development and Management of Regional Ocean Observing Systems Roger Pugliese briefed the Panel members on the development and management of Regional Ocean Observing Systems. Roger noted we are fortunate to be in a region that is being used as a test case for regional ocean observing systems. These will be designed to allow us to understand the ocean in a three-dimensional framework. Roger noted we have begun building links to these systems on the Ecosystem webpage and connections through the Internet Mapping Server. He noted that the systems will further our understanding of current and other factors, and help us and fishermen understand what is going on beneath the surface. He noted there will be some additional direct work to build the presentation of the data collected by these systems to support the FEP. Roger noted that we can go to the web site and view the presentations on the different systems, from the link to our Research and Monitoring Workshop. He indicated that fisheries has just jumped into this discussion in the last year or so. An opportunity was provided for us to be on the front end of this process. One NOS proposal dealt with ecological modeling, to assess the year-class strength of gag, based on temperature, and this would have fed
- 21 - Overview and Recommendations - Joint Habitat and Coral AP Meeting June 7-9, 2006 directly into the SEDAR process. The bottom line is that this would have been an excellent collaboration between oceanographers, fishery biologists and managers, but it didn’t get funded. Roger noted that we need to find the dollars to support feeding the information from oceanography directly into fisheries assessment and management.
The Advisory Panels support such collaboration and the funding it requires.
14) National Habitat Plan and Southeast Aquatic Resources Partnership Roger Pugliese addressed the National Fish Habitat Plan that is moving forward. He noted that it was initiated to a large degree by freshwater interests, to address fish habitat across the nation but also regionally. He noted that a partnership has been built in the SE, that covers a broader area than we normally consider. He noted the Council, states, USFWS, TNC, NMFS and many other organizations are participating. One of the first efforts coming out of the group is to develop a SE Aquatic Resource Plan. Roger noted that this will not reinvent the wheel, but will hopefully translate many of the recommendations from the FEP into action, beyond what the FEP could do by itself.
The Advisory Panels support these programs as partners in implementing the FEP.
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- 23 - SOUTH ATLANTIC FISHERY MANAGEMENT COUNCIL
ONE SOUTHPARK CIRCLE, SUITE 306
CHARLESTON, SOUTH CAROLINA 29407-4699
TEL 843/571-4366 or FAX 843/769-4520 Toll Free 1-866/SAFMC-10 E-mail: [email protected]
Web site: www.safmc.net
Louis Daniel, Chairman Robert K. Mahood, Executive Director George Geiger, Vice-Chairman Gregg T. Waugh, Deputy Executive Director
Overview & Recommendations
JOINT MEETING OF THE HABITAT ADVISORY PANEL AND CORAL ADVISORY PANEL
October 26-28, 2004
Francis Marion Hotel 387 King Street, Charleston, SC 29401
Issues addressed at this meeting included: 1) Ecosystem Pilot Projects and Action Plan for the Evolution of the Habitat Plan into a Fishery Ecosystem Plan for the South Atlantic Region; 2) The SAFMC Comprehensive Habitat and Fishery Ecosystem Plan Page and Coral and Benthic Habitats Internet Mapping Server; 3) Research Efforts on the Charleston Bump Essential Fish Habitat- Habitat Area of Particular Concern; 4) Deepwater Coral Habitat Research and Recommendations for Protection; 5) The Draft Oculina Closed Area Research and Monitoring Plan; 6) Gas Pipeline Development: Assessing Impacts on Nearshore and Deepwater Coral Habitats, Coordinated Council and NOAA Fisheries Comments, and Implications for Energy Policy Refinement; 7) Potential Actions for Comprehensive Ecosystem Amendment; and 8) Illegal Harvest of Living Coral.
1) Ecosystem Pilot Projects and Action Plan for the Evolution of the Habitat Plan into a Fishery Ecosystem Plan for the South Atlantic Region: With the Habitat Plan as a cornerstone, the Council is developing an ecosystem-based approach to resource management. Evolution of the Habitat Plan into a Fishery Ecosystem Plan (FEP), and transition from single species management to ecosystem-based management, will require a greater understanding of the South Atlantic Bight ecosystem and the complex relationships among humans, marine life and essential fish habitat. This effort will provide a more comprehensive understanding of the biological, social and economic impacts of management. The Habitat Plan will serve as the basis for the FEP. Updated life history and stock status information on managed species and the characteristics of the food web they exist within will be incorporated as well as social and economic research needed to fully address ecosystem-based management. Writing Teams (composed of AP members, experts from state and federal agencies, universities and Council staff) will review, update and expand existing chapters of the Habitat Plan and incorporate this material into new chapters for the FEP (e.g., Ecosystem Modeling and Research Needs to support Ecosystem-Based Management). The major areas to be addressed in the developing FEP and Comprehensive Ecosystem Amendment include the following: 1. Define the geographical boundaries of the ecosystem, including characterization of its biological, chemical and physical dynamics; 1
Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004
2. Assess ecological, human and institutional elements of the ecosystem: 3. Develop a conceptual model of the food web; 4. Describe the habitat needs of different life history stages for all managed species (including protected resources); 5. Calculate and characterize total removals (i.e., landings, effort, catch location, discards, and bycatch); 6. Develop indices of ecosystem health (e.g., biological indicators): 7. Establish long-term monitoring; and 8. Develop appropriate management including catch limits, gear regulations, zoning, etc.
5-Year System-Wide Evaluation
Habitat Plan Direction to Species Committees
Describe Ecosystem FISHERY ECOSYSTEM Option 1. Snapper PLAN Cumulative Impacts Grouper (SOURCE DOCUMENT) Amendment (13B) Summarize available data. Covers all FMPs & SAFE Reports - NOAA Species. Complete Initial Option 2. Snapper Fisheries provides Plan in 2005. Grouper (13B) & Comprehensive Stock Updated once every 5 Mackerel (16) Assessment and Fishery years. Amendment Evaluation Reports for all FMPs - updated annually or every 5 years Option 3. Comprehensive Amendment to Multiple or All Fishery Management Plans Biological, Social, Economic, Chemical/Physical, MMPA, & ESA/PR Information
Draft Timeline 2004/2005
Appoint Writing Teams from Habitat/Coral Advisory Panels, State and Draft Fishery Ecosystem Plan Federal Scientists, Universities & Staff (FEP) Draft Document Late 2005 (2004 & 2005)
Ecosystem-Based Workshops Management Draft Comprehensive (July’04-early 2005) Committee Meetings Amendment/EIS 2004 & 2005 Late 2005/Early 2006 1. Compliance with EFH Final Rule 2. Additional Coral HAPCs 3. Other Measures as necessary
2) Habitat/Ecosystem Page and Coral and Benthic Habitats Arc Internet Mapping System(IMS): Tina Udouj with the Florida Wildlife and Research Institute (FWRI) presented a summary of the developing Comprehensive Habitat and Fishery Ecosystem Plan web site and 2 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004
Internet Mapping Server (IMS). Panel members were requested to provide comments and recommendations that will aid in the further refinement of the Ecosystem site and IMS to better support regional ecosystem management.
Additional Background: The South Atlantic Council and the Florida Wildlife Research Institute (FWRI) partnered to develop a Comprehensive Habitat and Fishery Ecosystem Plan page that is accessible from the South Atlantic Council’s web site. FWRI is hosting an Internet Map Server (IMS) application with links to bottom type data that can be downloaded, associated metadata, substantial program information for the Council and links to related sites. The Web site is operated and maintained at FWRI in partnership with the South Atlantic Council. The Internet Map Server (IMS) component of this project brings the power of Geographic Information Systems (GIS) technology and Image Analysis tools to ordinary Internet browsers. The Coral and Benthic Habitats IMS will be an effective tool for displaying, sharing and querying coral, habitat and ecosystem information across the South Atlantic region. Researchers have a unique opportunity to access video and still imagery archives of coral and benthic habitats served from this site.
3) Research Efforts on the Charleston Bump Deepwater Essential Fish Habitat –Habitat Area of Particular Concern (EFH-HAPC): In 1998 the Charleston Bump was designated an EFH-HAPC for a number of managed species as part of the South Atlantic Council Habitat Plan and Comprehensive Habitat Amendment implementing Essential Fish Habitat in the South Atlantic Region. Over the last number of years a great deal more research has been conducted in the area including more extensive mapping and characterization of this unique system. Dr. George Sedberry, with the South Carolina Department of Natural Resources provided an update on research conducted at the Charleston Bump EFH-HAPC.
4) Deepwater Coral Habitat Research and Protection: Dr. Steve Ross with the University of North Carolina at Wilmington (UNCW) and John Reed with Harbor Branch Oceanographic Institute (HBOI) made presentations on research conducted on deepwater coral distribution and characterization in the South Atlantic Region. Andy Shepard, Director of the UNCW/NURC was contracted to coordinate the preparation of the reports prepared for the Council. The presentations encompassed research conducted to date and presented in the following reports developed for the Council:
GENERAL DESCRIPTION OF DISTRIBUTION, HABITAT, AND ASSOCIATED FAUNA OF DEEP WATER CORAL REEFS ON THE NORTH CAROLINA CONTINENTAL SLOPE (Ross, 2004); and DEEP-WATER CORAL REEFS OF FLORIDA, GEORGIA AND SOUTH CAROLINA: A SUMMARY OF THE DISTRIBUTION, HABITAT, AND ASSOCIATED FAUNA (Reed, 2004).
Council staff provided an overview of the integration of the establishment of new deepwater coral HAPCs into the Fishery Ecosystem Plan and Comprehensive Ecosystem Amendment development process. Panel members discussed the information provided to further refine previous recommendations on the establishment of new deepwater coral HAPCs in the South Atlantic Region. In addition, Council staff provided an overview of the preliminary development of a deepwater coral research and monitoring plan.
JOINT HABITAT AND CORAL AP CONCENSUS RECOMMENDATIONS The Advisory Panels unanimously recommend that the following sites be designated as deepwater coral HAPCs. Other possible sites off North Carolina identified by Dr. Ross
3 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004 should be considered for designation at a later date pending the acquisition of additional data. In addition, an area of recently identified Oculina occurs between the existing two satellite coral HAPCs and should be considered when adequate information becomes available.
Regulations in proposed deepwater Coral HAPCs: Recommended management measures in all the deepwater coral HAPC sites include the following: A) prohibit the use of bottom longline, bottom trawl, dredge, pot or trap; B) prohibit anchoring, the use of an anchor and chain or grapple and chain by any fishing vessel; C) prohibit fishing or possession of shrimp from the area; D) prohibit the possession of all species regulated by the coral FMP.
Clarification: To prohibit the collection of gorgonians in coral HAPCs (this would prevent any allowable harvest presently permitted under the coral plan, however, biomedical or taxonomic collections could occur with appropriate permits). The Council should investigate avenues to prohibit anchoring by all vessels. The Panels identified a future research priority to determine if damage is associated with other gears that may come in contact with the bottom (e.g., planers and downrigger weights).
Future considerations in FEP development: Explore the possibility for managing the deepwater corals and habitats that are not presently covered under some existing regulatory mechanism.
(Note: Dr. Bob George provided a letter to Doug Rader with possible deepwater Lophelia sites. It was the consensus of the Panels that the areas are likely already covered or are not adequately documented. If additional detailed information on new areas is provided, the areas could be considered in future reviews for possible coral HAPCs.)
A) Proposed Cape Lookout Lophelia Bank HAPC and B) Proposed Cape Fear Lophelia Bank HAPC (summarized from Ross, 2004) The Cape Lookout and Cape Fear Lophelia Banks areas are very rugged with some banks covering a vertical rise of 50-80 m in a distance of a kilometer or less with the most common in a depth range of 350-450 m. These features are very abrupt and may function as mini-seamounts. The banks occur in areas of high currents, and create eddies that retain nutrients and larvae. The Lophelia Banks off North Carolina are relatively isolated from the surrounding areas and are composed of almost pure Lophelia pertusa. It is often the only species present, although there are a couple of other hard coral species. The invertebrate community is quite diverse with Galatheid crabs appearing dominant and Echinoderms being common. Sea stars and brittle stars are common, including Novodinia antillensis, and Ophiacantha bidentata (brittle star). The brittle star, at the end of its range, is incredibly abundant. Several sea urchin species are present. The squat lobster, Eumunida picta is common. Other decapods include Bathynectes longispina, and Rochinia crassa, a spider crab. The fish species include Idiastion kyphos (a deepwater scorpaenid, known from only five or six specimens collected in the Caribbean), Scyliorhinus meadi (a cat shark), wreckfish, Polyprion americanus, and Physiculus karrerae (collected and observed a number of times). While a lot of data has been gathered over the last four years the coverage is still restricted. The cat shark they have observed has been seen several times, tightly associated with coral habitat. Some of these species, thought to be rare, may in fact just be associated with a specific habitat type. Other fish included Lophiodes beroe (a goosefish), Beryx decadactylus (red bream), Helicolenus dactylopterus (black-bellied rosefish) and Conger oceanicus. Red bream may be present in
4 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004 commercially fishable quantities. The rosefish and conger eel also may have commercial potential. Laemonema melanurum (coral hake), Laemonema barbatulum, Nzumia aequalis (a rattail), and Hoplostephus occidentalis (a roughy). Some of these species may have some commercial potential and it is uncertain if these species might be using the coral mounds as spawning habitats. Myctophid species occur in association with the reefs in dense numbers, and certainly can serve to transfer energy from the surface layers where they travel at night, to the bottom layers. Researchers indicate the canyon system of the Point, off Cape Hatteras, hosts a very different species composition than the Lophelia Banks. The deep coral reefs seem to have similar habitat functions as shallow coral reefs (increased faunal diversity, concentrated foods, obligate fauna). There seems to be a primary group of fishes unique to these habitats. Threats to these habitats include degradation of the coral habitat itself from fishing activity, energy exploration, cable laying, the over exploitation of coral dependent species, and others. The three Lophelia banks off North Carolina are the northernmost know on the Atlantic Coast until reaching Canada. The North Carolina banks are similar to each other but differ from those on the Blake Plateau. The fish community seems tightly coupled to the corals.
Threats: Researchers have noted that we should learn from the Oculina Bank and move rapidly to protect these diverse unique deepwater coral habitats. In addition, European deepwater corals have suffered extensive damage from trawling activities and the technology while not extensively used in the South Atlantic to fish deep systems exists. In addition, Panel members are very concerned over potential delay in implementing regulations once the word is out about these new habitats. In developing the regulations to protect the HAPCs fishing and non-fishing threats, including potential ones should be analyzed for each site.
Research Needs: Characterize and map the extent and distribution of the deepwater coral habitat, determine which biological resources are unique to or are dependant on deep coral bank habitat, determine the status of these complex habitats and their associated communities, including the origin of dead coral.
Future Action: Other undocumented Lophelia sites exist off North Carolina and future investigations should be conducted so if additional deepwater coral habitat exists, they can be considered for coral HAPC designation.
C) Proposed Stetson Reef Coral HAPC (summarized from Reed, 2004) The Proposed Stetson Reef Coral HAPC is on the outer eastern edge of the Blake Plateau, approximately 120 nm SE of Charleston, South Carolina, at depths of 640-869 m. Over 200 coral mounds up to 146 m in height occur over this 6174 km2 area that was first described by Thomas Stetson from echo soundings and bottom dredges. These were described as steep-sloped structures with active growth on top of the banks. Live coral colonies up to 50 cm in diameter were observed with a camera sled. Enallopsammia profunda was the dominant species in all areas although Lophelia pertusa was concentrated on top of the mounds. Densest coral growth occurred along an escarpment in the Region. Stetson reported an abundance of hydroids, alcyonaceans, echinoderms, actiniaria, and ophiuroids, but a rarity of large mollusks. Detailed geological maps of this region also indicate numerous coral mounds. Recent fathometer transects indicated dozens and possibly hundreds of individual pinnacles and mounds within the small surveyed region, which is only a fraction of the Stetson Bank area. From our fathometer transects, two pinnacle regions were selected. Three submersible dives were made on “Pinnacle 3” and four dives on “Stetson’s Peak” which is described below. A small subset of the Stetson Bank area was first mapped during six fathometer transects covering approximately 28 nm2, in which six major peaks or pinnacles and
5 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004 four major scarps were plotted. The base depth of these pinnacles ranged from 689 m to 643 m, with relief of 46 to 102 m. A subset of this was further mapped with 70 fathometer transects spaced 250 m apart (recording depth, latitude and longitude ~ every 3 seconds), covering an area of 1 x 1.5 nm, resulting in a 3-D bathymetric GIS Arcview map of a major feature, which we named Stetson’s Pinnacle. Stetson’s Pinnacle was 780 m at the south base and the peak was 627 m. This represents one of the tallest Lophelia coral lithoherms known, nearly 153 m in relief. The ridges were covered with nearly 100% Lophelia coral rubble, 15-30 cm colonies of live Lophelia, and standing dead colonies of Lophelia, 30-60 cm tall. Very little rock was exposed, except on the steeper exposed, eroded faces of the ridges. Towards the top of the peak, colonies of Lophelia, 30- 60 cm tall, were more common, and some rock ledges had nearly 100% cover of live Lophelia thickets. The top edge of the pinnacle was a 30 cm thick rock crust which was undercut from erosion; below this was a 90o escarpment of 3-6 m. The peak was a flat rock plateau at 625- 628 m and was approximately 0.1 nm across on a S-N submersible transect. Dominant sessile macrofauna consisted of scleractinia, stylasterine hydrocorals, gorgonacea and sponges. The colonial scleractinia were dominated by colonies of Lophelia pertusa (30-60 cm tall) and Enallopsammia profunda, and Solenosmilia variabilis were present. Small stylasterine corals (15 cm tall) were common and numerous species of solitary cup corals were abundant. Dominant octocorallia consisted of colonies of Primnoidae (15-30 cm tall), paramuriceids (60-90 cm), Isididae bamboo coral (15-60 cm), stolonifera, and stalked Nephtheidae (5-10 cm). Dominant sponges consisted of Pachastrellidae (25 cm fingers and 25- 50 cm plates), Corallistidae (10 cm cups), Hexactinellida glass sponges (30 cm vase), Geodia sp. (15-50 cm spherical), and Leiodermatium sp. (50 cm frilly plates). Although motile fauna were not targeted, some dominant groups were noted. No large decapods crustaceans were common although some red portunids were observed. Two species of echinoids were common, one white urchin and one stylocidaroid. No holothurians or asteroids were noted. Dense populations of Ophiuroidea were visible in close-up video of coral clusters and sponges. No large Mollusca were noted except for some squid. Fish consisted mostly of benthic gadids and rattails. On the steeper upper flank, from 671 to 625 m the density, diversity, and size of sponges increased; 15- 50 cm macro sponges were more abundant. Massive Spongosorites sp. were common, Pachastrellidae tube sponges were abundant, and Hexactinellida glass sponges were also common. On the peak plateau the dominant macrofauna were colonies of Lophelia pertusa (30- 60 cm tall), coral rubble, Phakellia sp. fan sponges (30-50 cm), and numerous other demosponges were abundant. No large fish were seen on top.
D) Proposed Savannah Lithoherms and East Florida Lophelia Reefs Deepwater Coral HAPC (summarized from Reed, 2004) The Savannah Lithoherms lie to the west, on the western Blake Plateau. It is shallower, 490-550m, deeper than the North Carolina sites. The currents are strong due to the proximity of the Gulf Stream. The Savannah Lithoherms include coral debris, coral thickets including octocorals and sponges. The thicket communities usually are on the edges of the ridges with a lot of standing dead coral which provides habitat for many other species. The system is extremely diverse with multiple trophic levels representing a whole ecosystem. The taxonomy for many species is being worked up now and includes a new species of gorgonian. Bushes of black coral and bamboo coral which both have commercial value are present. Researchers are concerned that as soon as this information is published someone may try to harvest it.
The proposed East Florida Lophelia Reefs HAPC is located at the foot of the Florida-Hatteras slope, near the Georgia border to as far south as Jupiter, Florida. Depth is 700-850 m, about 2500 feet, dropping into the Florida Straits. Adjacent muddy slopes are primary habitat for golden tilefish. Once off the muddy slopes there are a whole string of pinnacles. The bathymetry in the area includes hundreds of pinnacles with peaks from 25 to hundreds of feet with an average height of 100 feet. Due to the low resolution of existing charts many of the pinnacles are presently
6 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004 undocumented and unknown to fishermen. In May 2004, a cruise was conducted off the east coast of Florida to investigate purposefully selected unique bathymetry; every area visited was a Lophelia reef, even ones identified on fishing charts. The extent of these Lophelia reefs emphasizes the need direct effort to map these unique areas. The 300 newly-discovered Lophelia reefs and lithoherms are from a quarter to half-mile in diameter, with an average height of 100 feet, up to 500 feet with all coral and hard bottoms occupied by living organisms.
E) Proposed Miami Terrace Deepwater Coral HAPC (summarized from Reed, 2004) The Miami Terrace Escarpment reefs habitat includes gorgonians and all the hard corals. Many of the organisms may contain cancer fighting compounds which could be isolated and synthesized. The Miami Terrace is a 65-km long carbonate platform that lies between Boca Raton and South Miami at depths of 200-400 m in the northern Straits of Florida. It consists of high-relief Tertiary limestone ridges, scarps and slabs that provide extensive hard bottom habitat. At the eastern edge of the Terrace, a high-relief, phosphoritic limestone escarpment of Miocene age with relief of up to 90 m at depths of 365 m is capped with Lophelia pertusa coral, stylasterine hydrocoral (Stylasteridae), bamboo coral (Isididae), and various sponges and octocorals. Dense aggregations of 50-100 wreckfish were observed here during submersible dives in May 2004. Previous studies in this region include geological studies on the Miami Terrace and dredge- and trawl-based faunal surveys in the 1970s primarily by the University of Miami. Lophelia mounds are also present at the base of the escarpment (~670 m) within the axis of the Straits of Florida, but little is known of their distribution, abundance or associated fauna. Researchers have found thickets of Lophelia, Enallopsammia, and Madepora growing on elongate depressions, sand ridges and mounds. Large quantities of L. pertusa and E. profunda have also been dredged from 738-761 m. Recent submersible dives and fathometer transects at four sites indicated the outer rim of the Miami Terrace to consist of a double ridge with steep rocky escarpments. There were considerable differences among the sites in habitat and fauna; however, in general, the lower slopes of the ridges and the flat pavement on top of the terrace were relatively barren. However, the steep escarpments especially near the top of the ridges were rich in corals, octocorals, and sponges. Dominant sessile fauna consisted of the following Cnidaria: small (15- 30 cm) and large (60-90 cm) tall octocoral gorgonacea (Paramuricea spp., Placogorgia spp., Isididae bamboo coral); colonial scleractinia included scattered thickets of 30-60 cm tall Lophelia pertusa (varying from nearly 100% live to 100% dead), Madrepora oculata (40 cm), and Enallopsammia profunda; stylasterine hydrocorals (15-25 cm); and Antipatharia (30-60 cm tall). Diverse sponge populations of Hexactinellida and Demospongiae included: Heterotella sp., Spongosorites sp., Geodia sp., Vetulina sp., Leiodermatium sp., Petrosia sp., Raspailiidae, Choristida, Pachastrellidae, and Corallistidae. Other motile invertebrates included Asteroporpa sp. ophiuroids, Stylocidaris sp. urchins, Mollusca, Actiniaria, and Decapoda crustaceans (Chaceon fenneri and Galatheidae). Schools of ~50-100 wreckfish (Polyprion americanus), ~60-90 cm in length, were observed on several submersible dives along with blackbelly rosefish, skates, sharks, and dense schools of jacks.
F) Proposed Pourtales Terrace Deepwater Coral HAPC (summarized from Reed, 2004) The Pourtales Terrace, or “The Humps” as it is referred to by the local fishermen, is a shallower site, about 300 meters. The area is about ten miles due south of Alligator Reef. The whole terrace is comprised of Miocene-age rock with lots of pinnacles up to a hundred feet in height and a whole series of deepwater sinkholes that have not been described. One that has been investigated goes to 1800 feet in depth, and has sheer rock walls with abundant organisms. Researchers indicated that the bottom of one of these sinkholes is covered with dugong rib bones, verifying that manatees lived off Florida over ten million years ago. The principal fish seen there was snowy grouper, along with different eels. This was the only site where the snowy grouper was seen.
The Pourtalès Terrace provides extensive, high-relief, hard-bottom habitat, covering 3,429 km2
7 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004
(1,000 nm2) at depths of 200-450 m. The Terrace parallels the Florida Keys for 213 km and has a maximum width of 32 km. Reed et al. (2004a) surveyed several deep-water, high-relief, hardbottom sites including the Jordan and Marathon deep-water sinkholes on the outer edge of the Terrace, and five high-relief bioherms on its central eastern portion.. These submersible dives were the first to enter and explore any of these features. The upper sinkhole rims range from 175 to 461 m in depth and have a maximum relief of 180 m. The Jordan Sinkhole may be one of the deepest and largest sinkholes known. The high-relief area of the middle and eastern portion of the Pourtalès Terrace is a 55 km-long, northeasterly trending band of what appears to be karst topography that consists of depressions flanked by well defined knolls and ridges with maximum elevation of 91 m above the terrace. Further to the northeast of this knoll-depression zone is another zone of 40-m high topographic relief that lacks any regular pattern (Gomberg, 1976). The high-relief bioherms (the proposed HAPC sites within this region) lie in 198 to 319 m, with a maximum height of 120 m. A total of 26 fish taxa were identified from the sinkhole and bioherm sites. Species of potential commercial importance included tilefish, sharks, speckled hind, yellow-edge grouper, warsaw grouper, snowy grouper, blackbelly rosefish, red porgy, drum, scorpion fish, amberjack, and phycid hakes. Many different species of Cnidaria were recorded, including Antipatharia black corals, stylasterine hydrocorals, octocorals, and one colonial scleractinian (Solenosmilia variabilis).
The Tennessee and Alligator Humps are among dozens of lithoherms that lie in a region called “The Humps” by local fishers, ~14 nm south of the Florida Keys and south of Tennessee and Alligator Reefs. Three dives were made on Bioherm #3, approximately 8.5 nm NE of Bioherm#2. Bioherm #3 consisted of two peaks 1.05 nm apart with a maximum relief of 62 m. The North Peak’s minimum depth was 155 m and was 653 m wide at the base, which was 217 m deep at the east base and 183 m at the west side. The minimum depth of South Peak was 160 m and was about 678 m in width E to W at the base. The surrounding habitat adjacent to the mounds was flat sand with about 10% cover of rock pavement. From 213 m to the top, generally on the east flank of the mound, were a series of flat rock pavement terraces at depths of 210, 203, 198, 194, 183, and 171 m and the top plateau was at 165 m. Between each terrace a 30-45 degree slope consisted of either rock pavement or coarse sand and rubble. Below each terrace was a vertical scarp of 1-2 m where the sediment was eroded away leaving the edge of the terrace exposed as a horizontal, thin rock crust overhang of <1 m and 15-30 cm thick. The top of the bioherm was a broad plateau of rock pavement with 50-100% exposed rock, few ledges or outcrops, and coarse brown sand. Less time was spent on the western side, which was more exposed to the strong bottom currents. The west side of South Peak sloped more gradually than the eastern side, had more sediment, and no ledges were observed.
Fish Communities (summarized from Reed et al., 2004a) A total of 31 fish taxa, of which 24 were identified to species level, were identified from submersible videotapes and were associated with the deep-water sinkholes and high-relief bioherms. Few studies have directly documented deep-water fish associations with deep-water reef habitats in the western Atlantic. Most of the work has concentrated on the Charleston Bump region of the Blake Plateau off Georgia and South Carolina (Sedberry, 2001). Ross (pers. comm.) reported the following species are common to both the deep-water Lophelia reefs on the Blake Plateau off the Carolinas and those of this study: Chloropthalmus agassizi, Helicolenus dactylopterus, Hoplostethus sp., Laemonema melanurum, Nezumia sp., and Xiphias gladius. Species most common to the high-relief bioherms included deepbody boarfish, blueline tilefish, snowy grouper, and roughtongue bass. Some species were common at both the sinkhole and bioherm sites and included snowy grouper, blackbelly rosefish, and mora. In addition to the moribund swordfish observed in the Jordan Sinkhole, a swordfish was observed from the NR-1 submersible on top of Pourtales Terrace. Species of potential commercial importance included tilefish, sharks, speckled hind, yellowedge grouper, warsaw grouper, snowy grouper, blackbelly
8 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004 rosefish, red porgy, drum, scorpionfish, amberjack, and phycid hakes. Any of the features, both sinkholes and bioherms, could be overfished very easily since only a few individuals of the larger grouper species were present at any one site.
Benthic Communities (summarized from Reed et al., 2004a) The benthos at the bioherm sites was dominated by sponges, octocorals and stylasterids. A total of 21 taxa of Cnidaria were sampled or observed and 16 were identified to species level. These included 3 species of antipatharian black coral, 5 stylasterid hydrocorals, 11 octocorals with one possible new species, and 1 scleractinian (Solenosmilia variabilis). Eight species were associated only with the Pourtalès sinkholes and not the bioherms; these included two species of antipatharians; the octocorals Paramuricea placomus, Plumarella pourtalesii, Trachimuricea hirta; and the scleractinian Solenosmilia variabilis. Although Gomberg (1976) found evidence of skeletal remains of the colonial scleractinians Lophelia and Madrepora in sediment samples from the terrace, we did not see any colonies at our dive sites. Sponges identified from collections included 28 taxa. Five species of stylasterine hydrocorals were Distichopora foliacea, Pliobothrus echinatus, Stylaster erubescens, S. filogranus, and S. miniatus. On the flat pavement adjacent to the base of the mounds, stylasterids and antipatharian black coral bushes were common along with sea urchins and sea stars. The densities of sponges, stylasterid hydrocorals and octocorals were very high, especially on the plateaus and terraces of the bioherms on the Pourtalès Terrace. Maximum densities of sponges (>5 cm) on the plateaus ranged from 1 to 80 colonies per sq. m. Stylasterid coral densities ranged from 9 to 96 colonies and octocorals from 16 to 48 per sq. m. Densities of sponges and stylasterids also dominated the terraces and slopes of the bioherm sites but generally in lower densities than the peak plateaus whereas the octocorals generally had higher densities on the flanks (1-80 colonies per sq. m).
5) Oculina Closed Area Research and Monitoring Plan: Council staff provided an overview of the draft Oculina Closed Area Research and Monitoring Plan. This plan is part of a developing overall Evaluation plan for the Oculina Closed Area which encompasses Research and Monitoring, Outreach and Law Enforcement. Panel members were requested to provide comments and recommendations that will aid staff in the completion of the draft plan.
JOINT HABITAT AND CORAL AP CONCENSUS RECOMMENDATIONS: A) The Council should call this a management plan; B) We reiterate the need for enforcement; C) The Panels support maintaining a strong outreach and education component; D) The first priority research need is mapping and characterization of the habitats; and E) The second priority research need is to understand the role of the site relative to
9 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004 generating fish in surrounding areas (including spawning areas, movements around the area, and identifying stocks around the site).
6) Gas Pipeline Development: Assessing Impacts on Near-shore and Deepwater Coral Habitats: Myra Brouwer with Council staff made a presentation on gas pipeline development, the coordination of Council and NOAA Fisheries comments, and the implications for Energy Policy refinement. Douglas Rader addressed the coordinated revision process for the Habitat Policy Statement on Energy Development to address gas pipeline development as well as other emerging energy processes.
JOINT HABITAT AND CORAL AP CONCENSUS RECOMMENDATIONS: Revise the Energy Policy through a coordinated effort including Council staff, Jocelyn Karazsia, NOAA Fisheries Habitat Conservation Division, a recently hired intern with Environmental Defense and the Habitat and Coral Advisory Panels. Additional issues for consideration include identification of proposed deepwater coral HAPCs and the use of conservation easements to generate funding streams for underwriting needed management and research.
7) Fishery Ecosystem Plan and Comprehensive Ecosystem Amendment Development: Panel members were requested to provide comments on the potential list of actions for consideration in a developing Comprehensive Ecosystem Amendment. Discussions revolved around a draft list of proposed measures for the Amendment.
SUMMARY OF POTENTIAL MANAGEMENT MEASURES FOR THE SAFMC COMPREHENSIVE ECOSYSTEM AMENDMENT:
CALCULATE AND CHARACTERIZE TOTAL REMOVALS (i.e., landings, effort, catch location, gear type/usage, discards, and bycatch including marine mammals and birds) 1. Identify all users: A. Require a permit to fish for, harvest, or possess any EEZ resource for all: (i) Commercial vessels (includes commercial and for-hire) (ii) Private recreational anglers
2. Calculate and characterize removals: A. Continue to implement the Atlantic Coastal Cooperative Statistics Program – catch and effort (trip tickets, logbooks & MRFSS); discards, bycatch and protected resources; socio- economic….. If a permit is required for private recreational anglers, the methods used to calculate catch and removals from anglers could be modified to sample from a known universe (permit holders). This alternative assumes completion of work to link the permits and logbook/landings databases in the southeast.
3. Compliance with the Essential Fish Habitat (EFH) final rule: A. Refine Essential Fish Habitat (EFH) and Essential Fish Habitat-Habitat Areas of Particular Concern (EFH-HAPCs) designations.
B. Identify new Essential Fish Habitat (EFH) and Essential Fish Habitat-Habitat Areas of Particular Concern (EFH-HAPCs) as necessary.
10 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004
C. Address measures to reduce impacts of fishing and non-fishing impacts on Essential Fish Habitat (EFH) and Essential Fish Habitat-Habitat Areas of Particular Concern (EFH-HAPCs) as necessary. (i) Establish “Allowable Trawling Areas”
4. Coral, Coral Reef and Live/Hardbottom Habitat Fishery Management Plan: A. Establish additional Coral Habitat Areas of Particular Concern (HAPCs) – gear prohibitions from existing HAPCs would apply within these new HAPCs. B. No action
5. Enforcement/Data Collection/Safety at Sea: A. Require VMS on all commercial, for-hire and private recreational vessels. Private recreational vessels could use a “chip” that is imbedded in existing electronic gear or some type of acoustic monitoring. B. Require VMS on all commercial and for-hire vessels. C. Require VMS on all commercial vessels.
6. Changes to other Fishery Management Plans A. Snapper Grouper - establish management units and move to unit-based management: (i) Red porgy - allow increase in harvest (rebuilding) (ii) Black sea bass - reduce catches (overfishing/overfished) (iii) Vermilion snapper - reduce catches (overfishing) (iv) Snowy grouper - reduce catches (overfishing/overfished) (v) Golden tilefish – reduce catches (overfishing)
B. Mackerel (i) Atlantic king mackerel - reduce TAC to address expected effort shift from snapper grouper regulations.
C. Protected Species Interactions: (i) Recommendations to Take Reduction Teams (ii) Recommendations to States (iii) Recommendations to ASMFC
JOINT HABITAT AND CORAL AP CONCENSUS RECOMMENDATIONS: Comprehensive Amendment Measures: the FEP provide the foundation to develop an allowable trawling area for the deepwater trawl fishery noting adequate information should be available to define the fishing area from the VMS system required for the rock shrimp fishery. This measure could enhance protection of unique habitat values of deepwater coral/habitat including the proposed deepwater coral HAPCs and deepwater EFH-HAPCs including the Charleston Bump EFH-HAPC.
Suggested the long-term implementation of Ecosystem Management include developing allowable gear zones considering gear use and location of historic fisheries.
Fishery Ecosystem Plan State Breakout Session Summaries: Break-out sessions of the four state habitat sub-panels (coral AP members selected which panel discussion to join) brainstormed on what can be included in the FEP for habitat protection. Panels were asked to come up with two lists: one for inclusion in the plan next fall and the other to be implemented in the long-term.
11 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004
North Carolina Sub-panel breakout session: Short-term list for draft plan: • Add reverse osmosis discharges to the list of threats, especially where the discharges are in freshwater. • The big push for development of rural areas in NC is being done under nationwide and other general permits, and therefore undergoes no review causing general environmental degradation in Primary Nursery Areas. • A cumulative impacts analysis and an assessment of the impacts of coastal development on fishery habitat quality should be included. • Incorporate some of the NC areas of Lophelia identified by Steve Ross into the list of proposed MPAs, or some other protective mechanism. • The plan should designate nursery areas for Council species. • Include as much information in the plan as possible that maps the location of effort expended, since that will help to highlight habitats used by the species. • Model the impacts of coastal development on the fisheries, i.e., highlight the connection between habitat degradation, and the loss of fishery productivity, noting that all the management measures in the world will be ineffective, unless habitat quantity and quality are maintained. • Include lists of researchers who are doing habitat work, as well as the kind of work they are doing, to facilitate identification of work that can be directly linked to development of better ways to protect fisheries and habitat sustainability. This would identify current areas of research that we can use to evaluate management measures. • Specify, to the extent we can do so, the necessary freshwater inflows for sustaining spawning and nursery habitats, as well as nearshore ocean habitats. This might be a long- term need, but there is some information that we might be able to include now. • Support the development of indicators of ecosystem health that can be used throughout the Council’s jurisdiction. Long-term items: • Develop habitat-production relationships for all species, especially for species associated with deepwater corals. We noted that this will take a long time to develop, but would provide a better science basis for convincing society that adverse impacts should be avoided. • Identify those habitats for which mapping is incomplete or non-existent, and identify resources and strategies for obtaining the information. This should include mapping of subtidal oyster bed habitats, and Submerged Aquatic Vegetation habitats.
South Carolina Sub-panel breakout session: • Recommend requiring all commercial and for-hire vessels (charter and headboats) have Vessel Monitoring Systems to identify area fished related to habitats. • The FEP better define ecosystems so we will have a better ability to prioritize how the Council will formulate management regulations. • Identify different uses of habitats in state versus federal waters, such as management areas, and research areas. • The plan should provide the vehicle for development of flexible or adaptive management.
Georgia Sub-panel breakout session: • Identify water flows, particularly in the Savannah and the challenge to define a desired outcome, ecologically.
12 Overview and Recommendations - Joint Habitat and Coral AP Meeting Oct. 26-28, 2004
• Concern over the proliferation of downstream coastal development, especially docks, as well as the development of more isolated areas, such as coastal hammocks. • Concern over manatees and discharges that might bring them into harms way. • The occurrence of exotic species, such as green mussels, and making sure their impact is included in the FEP. • Harbor impacts, and need for regional consideration given to such harborification. • The occurrence of disease relative in blue crabs and penaeid shrimp, and the changes in susceptibility as a consequence of runoff and need to allocate funds to deal with this issue. • Possible additional EFH-HAPCs could include: cool water seeps that provide needed refugia for certain species; good quality, small watershed, coastal streams that might be relatively unimpaired, and serve as baseline systems Long-term items: • Include the impact of sea-level rise on the barrier islands and marsh systems in coastal GA. • Airborne deposition of mercury and other contaminants.
Florida Sub-panel breakout session: • The FEP needs to include headwaters to the bluewaters, in order to effectively deal with the ecosystem. All aspects have to be addressed for this to be a true ecosystem plan. • Human interactions aspects have to be included, including harbors, coastal development, beach nourishment, land use changes. • Water management operations, related to effluents, both the good and bad of it, contaminants, as well as nutrients. Both increases and reductions in discharges need to be considered. • Exotic species need to be included. • The food web needs to be considered, for the entire range of a species. • The whole universe of management measures should be evaluated in the assessment, not just particular ones. • Online reporting as a possibility for recreational fishermen. • Relative to calculating total removals, there needs to be some system in place for ensuring full disclosure of information.
8) Illegal Harvest of Living Coral: At the request of Council staff, David Dale, NOAA Fisheries, Habitat Conservation Division briefed the Panels on the issue of illegal harvest of live coral. He indicated that the cases investigated appear to involve Indo-Pacific corals in aquarium shops, or illegally-harvested live rock, or cultured live rock. He noted he had hoped to have information from the agent, but didn’t have the numbers yet. He will keep the Panels and Council posted on this issue. Jenny Wheaton, FWRI indicated that they do all the identifications for enforcement, and they haven’t seen any increase. If information can be provided to her, she will try to track down the extent. Terry Gibson noted that he felt the issue is rampant in South Florida and a case was recently made. Stephen Blair noted that nearshore small cases are harder to prosecute and follow.
JOINT HABITAT AND CORAL AP CONCENSUS RECOMMENDATIONS: The Panels agreed that the frequency and extent of this activity needs to be explored to determine if this issue is perception versus reality.
13 HABITAT AND FAUNA OF DEEP-WATER CORAL REEFS OFF THE SOUTHEASTERN USA
A Report to the South Atlantic Fishery Management Council Addendum to 2004 Report 2005-2006 Update- East Florida Reefs
by John K. Reed Harbor Branch Oceanographic Institution 5600 U.S. 1, North, Fort Pierce, FL 34946 Phone- 772-465-2400 x205, Fax- 772-461-2221 Email- [email protected]
Contract No: SA-05-09-FL/FWRI Submitted to: South Atlantic Fishery Management Council One Southpark Circle, Suite 306 Charleston, SC 29407
[All rights reserved. Authorization requested by the author for photocopying or electronic distribution of any parts of this document. Copying or electronic distribution of tables or figures must include accompanying caption with complete citation.]
August 17, 2006
ABSTRACT
In 2004 a Summary Report (Reed 2004) was compiled by the PI at the request of the South Atlantic Fishery Management Council (SAFMC) to provide a preliminary, general summary on the status of current knowledge concerning deep-water (> 200 m) reefs off the southeastern U.S. from Florida to North Carolina. The purpose was to prioritize areas of deep-water, live-bottom habitats for: 1) potential designation as Habitat Areas of Particular Concern (HAPC) or Marine Protected Areas (MPA) by the SAFMC, and 2) high-resolution habitat maps and habitat characterization studies.
The following report is an update to the 2004 Report that provides new data collected from eight expeditions using submersible or ROV off eastern Florida during 2005 and 2006. Based on the 2004 Report and the data from this report that was presented by the PI to the Coral and Habitat Advisory Panels (SAFMC meeting, June 2006), the SAFMC has proposed six new deep-water reef HAPCs off southeastern US. The resource potential of the deep-water habitats in this region is unknown in terms of fisheries and novel compounds yet to be discovered from associated fauna that may be developed as pharmaceutical drugs. Activities involving bottom trawling, pipelines, or oil/gas production could negatively impact these reefs.
JUSTIFICATION
The South Atlantic Fishery Management Council (R. Pugliese) requested that this update to the 2004 Report on the state of knowledge of deep sea coral ecosystems (DSCE) off Florida be available in time for the Coral and Habitat Advisory Panels meeting of the SAFMC, June 9, 2006. The Council needs immediate scientific data and maps as it considers designation of new Habitat Areas of Particular Concern (HAPC) to protect DSCE areas. Such protection may be needed to prevent long-term (perhaps permanent) damage, such as has occurred on shallower Oculina reefs off Florida and Lophelia banks in the northeastern Atlantic, both destroyed in part by trawling. After trawlers were banned from the Oculina HAPC, there is justified concern that trawlers may move to deeper habitats in search of valuable commercial fisheries, such as royal red shrimp or benthic finfish. The resource potential of the deep-water habitats in this region is unknown in terms of fisheries and novel compounds yet to be discovered from associated fauna that may be developed as pharmaceutical drugs. Although these habitats are not currently designated as MPAs or HAPCs, they are incredibly diverse and irreplaceable resources. Activities involving bottom trawling, pipelines, or oil/gas production could negatively impact these reefs.
OBJECTIVES
Objectives of this report and accompanying DVD are the following: 1) Compile a list of research cruises that explored the deep-water reefs off eastern Florida from 2005 to 2006. 2) Compile list of submersible dives, including dive number, date, location, GPS coordinates, depth, and habitat type for each dive (DVD- Excel file).
2 3) Provide Powerpoint presentation of this report, including insitu digital still images and video of newly discovered bottom habitat, that was presented to the Coral and Habitat Advisory Panels meeting of the SAFMC, June 9, 2006 (DVD- ppt file). 4) Provide Cruise Report from the following expedition: Florida’s Deep-Water Oases: Exploration of Deep Reef Ecosystems, May 31- June 9, 2006 (DVD- ppt file). This expedition provided for the first time an assessment of the biodiversity and relative abundance of the benthic, fish and zooplankton communities; geological features; physical processes; and biochemical compounds of interest for drug discovery within a deep-water reef ecosystem.
INTRODUCTION
Deep-Sea Coral Reefs (from Reed, 2004) Deep-water reefs are sometimes defined as bioherms, coral banks, or lithoherms (Teichert, 1958; Stetson et al., 1962; Neumann et al., 1977; Wilson, 1979; Reed, 1980; Freiwald et al. 1997; Fosså et al. 2000; Paull et al., 2000). Some deep-water reefs consist of caps of living coral on mounds of unconsolidated mud and coral debris, such as some Oculina and Lophelia coral reefs (Reed 2002a,b; Reed et al. 2005, 2006), whereas deep-water lithoherms are defined as high-relief, lithified carbonate limestone mounds rather than unconsolidated mud mounds (Neumann et al., 1977). Rogers (1999) has suggested that deep-water coral bioherms fall within the definition of a coral reef based on their physical and biological characteristics. Various types of deep-water, high-relief bioherms are common off the southeastern United States, along the base of the Florida- Hatteras Slope, on the Blake Plateau, in the Straits of Florida, and eastern Gulf of Mexico (Reed et al., 2005, 2006). Only a small percentage of deep-water reefs have had their benthic and fish resources characterized.
Florida DSCE Deep sea coral ecosystems (DSCE) in U.S. EEZ waters exist along the eastern and southwest Florida shelf slope (in addition to the Oculina Marine Protected Area and deep shelf-edge reefs with hermatypic coral). These include a variety of high-relief, hard-bottom, live-bottom habitats at numerous sites along the base of the Florida-Hatteras Slope off northeastern and central eastern Florida, the Straits of Florida, the Miami Terrace and Pourtales Terrace off southeastern Florida, and the southwestern Florida shelf slope. The predominate coral on these reefs are the azooxanthellate, colonial scleractinian corals, Lophelia pertusa , Madrepora oculata , and Enallopsammia profunda ; various species of hydrocorals of the family Stylasteridae, and species of the bamboo octocoral of the family Isididae. Various types of high-relief, live-bottom habitat have been discovered in the area: Lophelia mud mounds, lithoherms, sinkholes, ancient Miocene escarpments and karst topographic features (Reed 2002b; Reed et al., 2004a,b, 2005, 2006). These all provide hard-bottom substrate and habitat for sessile macrofauna including deep-water corals, octocorals (gorgonians), black coral, and sponges, which in turn provide habitat and living space for a relatively unknown but biologically rich and diverse community of associated fish, crustaceans, mollusks, echinoderms, polychaete and sipunculan worms, and other macrofauna, many of which are undoubtedly undescribed species.
Recent research expeditions by Principal Investigator (PI), J. Reed, Harbor Branch Oceanographic Institution (HBOI), using HOVs (human occupied vehicle) and ROVs (remotely
3 operated vehicle) along with previous research by the PI in the 1990s and 1980s, have compiled new information on the status, distribution, habitat, and biodiversity of some of these relatively unknown and newly discovered deep reef ecosystems. In 2004, during a State of Florida funded mission with the Johnson-Sea-Link (JSL ) Submersible, the PI discovered nearly 300 potential targets during echosounder transects that may be newly discovered deep-water reefs off the east coast of Florida, some of which are up to 168 m (550 feet) in height at depths of 732 m (2400 feet) (Reed and Wright, 2004; Reed et al., 2004b, 2005, 2006). Expeditions in 2002 and 2003 for biomedical research by the PI and funded by the National Oceanic and Atmospheric Administration’s Office of Ocean Exploration (NOAA OE) enabled preliminary exploration of additional deep-water reef sites in the western Atlantic (Blake Plateau) and eastern Gulf of Mexico on southwest Florida shelf slope (Reed, 2003, 2004; Reed and Pomponi, 2002b; Reed et al., 2002, 2003, 2004d, 2006). These were the first HOV and ROV dives ever to document the habitat and benthic biodiversity of some of these relatively unknown deep-water reefs.
This report provides new information based on eight expeditions on deep-water reefs off eastern Florida and Straits of Florida using submersible or ROV during 2005 and 2006.
RESULTS
Cruise Summaries The following summarizes all expeditions that explored deep-water reefs off eastern Florida during 2005 and 2006.
1) Title : Florida’s Deep-Water Oases: Exploration of Deep Reef Ecosystems Institution : Harbor Branch Oceanographic Institution (HBOI), Division of Biomedical Marine Research (DBMR) Principal Investigators: John Reed, Amy Wright (HBOI) Ship/Submersible : R/V Seward Johnson , Johnson-Sea-Link I submersible Dates : April 4-15, 2005 Location : Bahamas- Bimini, Cay Sal; Florida- Miami Terrace Number of submersible dives : 18 New reef sites discovered : 2 new reef sites ground truthed w/ sub
2) Title : Center of Excellence for Biotechnology and Marine Biomedical Research- Exploration of Deep Reef Ecosystems Institutions : Harbor Branch Oceanographic Institution (HBOI), Division of Biomedical Marine Research (DBMR); Florida Atlantic University, Center of Excellence for Biotechnology and Marine Biomedical Research Principal Investigators : John Reed, Amy Wright, Shirley Pomponi (HBOI); Russ Kerr, Frank Mari (FAU) Ship/Submersible : R/V Seward Johnson , Johnson-Sea-Link I submersible Dates : August 2-16, 2005 Location : Miami Terrace, Straits of Florida Number of submersible dives : 27 New reef sites discovered : 9 new reef sites ground truthed w/ sub
4 3) Title : Florida’s Deep-Water Oases: Exploration of Deep Reef Ecosystems Institutions : Harbor Branch Oceanographic Institution (HBOI), Division of Biomedical Marine Research (DBMR); Oregon Institute of Marine Biology, NOVA Southeastern University, Smithsonian Institution, NOAA Office of Ocean Exploration (funding agency) Principal Investigators : Sandra Brooke (OIMB), John Reed (HBOI), Charles Messing (NOVA) Ship/Submersible : R/V Seward Johnson , Johnson-Sea-Link I submersible Dates : Nov. 7-20, 2005 Location : Florida Lophelia Reefs, Miami and Pourtales Terrace Number of submersible dives : 14 New reef sites discovered : 8 new reef sites ground truthed w/ sub; 31 new potential targets from echosounder
4) Title : Seafarer Proposed Natural Gas Pipeline Route Deep-Water Survey Institutions : Harbor Branch Oceanographic Institution (HBOI), ENSR Corp. Principal Investigator : John Reed (HBOI) Ship/Submersible : R/V Seward Johnson , Johnson-Sea-Link I submersible Dates : February 28- March 7, 2006 Location : Florida Lophelia Reefs, Straits of Florida Number of submersible dives: 9 New reef sites discovered : 1 new reef site ground truthed w/ sub; 18 nm ground truthed w/ sub
5) Title : Calypso Proposed Natural Gas Deep-Water Port Site Survey Institutions : NOVA Southeastern University, Harbor Branch Oceanographic Institution (HBOI), Florida Fish and Wildlife Research Institute, Naval Surface Warfare Center, Suez Inc. Principal Investigator : Charles Messing (NOVA), John Reed (HBOI), Sandra Brooke (FWRI) Ship/Submersible : R/V Walton Smith , TONGS ROV Dates : April 15-18, 2006 Location : Miami Terrace, Straits of Florida Number of ROV dives : 15 transect legs New reef sites discovered : 24 2 nm ground truthed w/ ROV; 36 reef/hard bottom sites recorded
6) Title : Calypso Proposed Natural Gas Pipeline Route Deep-Water Survey Institutions : NOVA Southeastern University, Harbor Branch Oceanographic Institution (HBOI), Naval Surface Warfare Center, Suez Inc. Principal Investigator : Charles Messing (NOVA), John Reed (HBOI) Ship/Submersible : R/V Walton Smith , TONGS ROV Dates : May 11-15, 2006 Location : Miami Terrace, Straits of Florida
5 Number of ROV dives : 15 transect legs New reef sites discovered : 50 nm ground truthed w/ ROV; 51 reef/hard bottom sites recorded
7) Title : Florida’s Deep-Water Oases: Exploration of Deep Reef Ecosystems Institution : Harbor Branch Oceanographic Institution (HBOI), Division of Biomedical Marine Research (DBMR); University of Miami, RSMAS Principal Investigators : John Reed, Shirley Pomponi, Amy Wright (HBOI); Mark Grasmueck, Gregor Eberli (UM) Ship/Submersible : R/V Seward Johnson , Johnson-Sea-Link II submersible Dates : May 22-30, 2006 Location : Bahamas- Bimini, Lucaya; Florida- Miami Terrace Number of submersible dives : 12 New reef sites discovered : 9 new reef sites ground truthed w/ sub; four 2x2 nm high def multibeam maps groundtruthed
8) Title : Florida’s Deep-Water Oases: Exploration of Deep Reef Ecosystems Institution : Harbor Branch Oceanographic Institution (HBOI), Division of Biomedical Marine Research (DBMR); University of Miami, RSMAS; Florida Fish and Wildlife Research Institute; NOVA Southeastern University; University of Florida Principal Investigators : John Reed, Tracey Sutton, Tammy Frank, Marsh Youngbluth (HBOI); Charles Messing (NOVA); Chuck Jacoby (UF); Robert Ginsburg, Chris Langdon (UM); Tina Udouj (FWRI) Ship/Submersible : R/V Seward Johnson , Johnson-Sea-Link II submersible Dates : May 31- June 9, 2006 Location : Miami Terrace, Straits of Florida Number of submersible dives : 16 New reef sites discovered : 2 reef sites ground truthed w/ sub. Detailed ecological assessment of the biodiversity and relative abundance of the benthic, fish and zooplankton communities; geological features; physical processes within this ecosystem; and biochemical compounds of interest for drug discovery.
REFERENCES
Reed, J.K. 2002a. Comparison of deep-water coral reefs and lithoherms off southeastern U.S.A. Hydrobiologia 471: 57-69.
Reed, J.K. 2002b. Deep-water Oculina coral reefs of Florida: biology, impacts, and management. Hydrobiologia 471: 43-55.
Reed, J.K. 2003. Deep-water coral reefs off Southeastern USA. National Geographic Society, Classroom Exploration of the Oceans, 2003, Keynote presentation [http://www.coexploration.org/ceo].
6 Reed, J.K. 2004a. General description of deep-water coral reefs of Florida, Georgia and South Carolina: A summary of current knowledge of the distribution, habitat, and associated fauna. A Report to the South Atlantic Fishery Management Council, NOAA, NMFS, 71 pp.
Reed, J.K. 2004b. Medicines from the deep sea: exploration of the Gulf of Mexico. The Slate, American Academy of Underwater Sciences, Vol. 1, 2004, p. 10-11.
Reed, J. K. and S. Pomponi. 1999. Submersible and scuba collections in the Gulf of Mexico, Florida Keys National Marine Sanctuary, and Florida Straits: Biomedical and biodiversity research of the benthic communities with emphasis on the porifera and gorgonacea, August 5-25, 1999.
Reed, J. K. and S. Pomponi. 2002a. Submersible and scuba collections in the Gulf of Mexico, Florida Keys National Marine Sanctuary, and Straits of Florida: biomedical and biodiversity research of the benthic communities with emphasis on Porifera and Gorgonacea. Final Cruise Report.
Reed, J.K. and S. Pomponi. 2002b. Islands in the Stream 2002: Exploring Underwater Oases. Mission Three: Summary. Discovery of new resources with pharmaceutical potential. Final Cruise Report.
Reed, J.K. and A. Wright. 2004. Final cruise report. Submersible and scuba collections on deep-water reefs off the east coast of Florida, including the Northern and Southern Straits of Florida and Florida Keys National Marine Sanctuary for biomedical and biodiversity research of the benthic communities with emphasis on the Porifera and Gorgonacea, May 20- June 2, 2004. Conducted by the Center of Excellence, HBOI and FAU, 54 pp.
Reed, J.K., R.H. Gore, L.E. Scotto, and K.A. Wilson. 1982. Community composition, structure, aereal and trophic relationships of decapods associated with shallow- and deep-water Oculina varicosa coral reefs. Bulletin of Marine Science 32: 761-786.
Reed, J.K., S. Pomponi, T. Frank, and E. Widder. 2002. Islands in the Stream 2002: Exploring Underwater Oases. Mission Three: Summary. Discovery of new resources with pharmaceutical potential; vision and bioluminescence in deep-sea benthos. NOAA Ocean Exploration web site: http://oceanexplorer.noaa.gov/explorations/02sab/logs/summary/summary.html , 29 pp.
Reed, J.K., A. Wright, and S. Pomponi. 2003. Discovery of new resources with pharmaceutical potential in the Gulf of Mexico. Mission Summary Report, 2003 National Oceanic and Atmospheric Administration Office of Ocean Exploration, 31 pp.
Reed, J. K., A, Wright, S. Pomponi. 2004. Medicines from the Deep Sea: Exploration of the Northeastern Gulf of Mexico. In: Proceedings of the American Academy of Underwater Sciences 23 th Annual Scientific Diving Symposium, March 12-13, 2004, Long Beach, California, p. 58-70.
7 Reed, J.K., S. Pomponi, A. Wright, D. Weaver, and C. Paull. 2005. Deep-water sinkholes and bioherms of South Florida and Pourtales Terrace- Habitat and Fauna. Bulletin of Marine Science 77:267-296
Reed, J.K., A. Shepard, C. Koenig, K. Scanlon, and G. Gilmore. 2005. Mapping, habitat characterization, and fish surveys of the deep-water Oculina coral reef Marine Protected Area: a review of historical and current research. Pp. 443-465, In (A. Freiwald, J. Roberts, Ed .), Cold- water Corals and Ecosystems, Proceedings of Second International Symposium on Deep Sea Corals, Sept. 9-12, 2003, Erlanger, Germany, Springer-Verlag, Berlin Heidelberg.
Reed, J.K., D. Weaver, S.A. Pomponi. 2006. Habitat and fauna of deep-water Lophelia pertusa coral reefs off the Southeastern USA: Blake Plateau, Straits of Florida, and Gulf of Mexico. Bulletin of Marine Science 78(2): 343-375.
Reed, J.K., C. Koenig, A. Shepard. 2006 (in review). Effects of bottom trawling on a deep- water Oculina coral ecosystem. Proceeding of 3 rd International Deep Sea Coral Symposium, Bulletin of Marine Science.
8
REVIEW OF DISTRIBUTION, HABITATS, AND ASSOCIATED FAUNA OF DEEP WATER CORAL REEFS ON THE SOUTHEASTERN UNITED STATES CONTINENTAL SLOPE (NORTH CAROLINA TO CAPE CANAVERAL, FL)
Steve W. Ross*
UNC-Wilmington, Center for Marine Science 5600 Marvin Moss Ln. Wilmington, NC 28409
*Currently assigned (through Intergovernmental Personnel Act) to: US Geological Survey, Center for Coastal & Watershed Studies, St. Petersburg, FL
Report Prepared for the South Atlantic Fishery Management Council One Southpark Circle, Suite 306 Charleston, SC 29407
16 May 2006 (second edition)
TABLE OF CONTENTS
INTRODUCTION 1
HISTORY OF DEEP CORAL RESEARCH IN THE SEUS 2
DEEP SEA CORALS OF THE SEUS 5
NORTH CAROLINA DEEP CORAL BANKS 6 Biological Communities 8 Cape Lookout Lophelia Bank A 8 Cape Lookout Lophelia Bank B 12 Cape Fear Lophelia Bank 20
CORAL BANKS OF THE BLAKE PLATEAU (SC to FL) 20
MAPPING DEEP CORAL BANKS 27
DEEP CORAL BANK FISH COMMUNITY DATA 27
SUMMARY AND RECOMMENDATIONS 32 Recommendations 32
ACKNOWLEDGMENTS 33
LITERATURE CITED 33
INTRODUCTION
Most habitats of the continental slope, and even the shelf edge, are poorly studied and in many cases completely unknown. These deeper areas, between 100 to 1000 m, are important frontiers, offering a transition from the continental shelf to the deep sea. Fisheries are expanding rapidly into these deep regions (Roberts 2002), and hydrocarbon exploration and development are now also exploiting these depths. Off the southeastern US (SEUS) coast (including the Gulf of Mexico) there are several unique and productive deep water habitats that have been difficult to study with conventional methods because the bottom topography is very rugged and the habitats are overlain by extreme currents (i.e., Gulf Stream). This report briefly summarizes data relevant to selected such poorly studied and vulnerable habitats (i.e., deep coral mounds) on the SEUS continental slope from Cape Lookout, NC to just south of Cape Canaveral, FL deeper than 200 m. Deep (or cold water) coral reef systems are receiving more attention worldwide. There is increasing evidence that deep water (aphotic) corals are important fish habitat (Costello et al. 2005), a repository of data on ocean climate and productivity (Adkins et al. 1998; Williams et al. in press), and are hotspots of increased biodiversity, including undescribed species. This is underscored by the growing literature and management concern directed toward these ecosystems (e.g., Morgan and Pizer 2005; Deep Sea Coral Habitat Act introduced in 2005). These habitats appear to be more extensive and important than previously known (e.g., SGCOR 2004; S.W. Ross, unpubl. data), while at the same time being severely threatened by a variety of activities (e.g., fishing, energy exploration) (Rogers 1999; Koslow et al. 2000). Although more extensive surveys are needed, Lophelia reefs (plus many other coral species) appear to populate the SEUS continental slope in great abundance (Stetson et al. 1962; Paull et al. 2000; Reed 2002a; Popenoe and Manheim 2001; P. Popenoe, pers. comm.). By one estimate the SEUS and Gulf of Mexico have the most extensive deep coral areas in the US (Hain and Corcoran 2004); however, these large regions are poorly explored (even considering recent expeditions). These high profile features concentrate exploitable resources and enhance local productivity in ways similar to seamounts (Rogers 1994; Koslow 1997), but this has not been adequately examined. For these reasons, locating, describing, and mapping deep corals and conducting basic biological studies in these habitats are considered global priorities (McDonough and Puglise 2003; Roberts and Hirshfield 2003; Puglise et al. 2005). Deep water coral habitat may be more important to western Atlantic slope species than previously known. Commercially-exploited deep-water species congregate around deep coral habitat, and evidence of fishing activities (trash, lost gear) was observed on some deep coral banks. Various invertebrates, particularly galatheid squat lobsters and echinoderms, are abundant on these deep reefs, playing roles of both predator on and food for the fishes. The deep reefs are oases offering both shelter and food. Additionally, the coral thickets are surrounded by extensive coral rubble habitat which preliminary data indicate also support a diverse fauna. Deep coral reefs of all types have been poorly studied, particularly so in the western Atlantic. References on deep coral banks within the SEUS EEZ are largely geological with a few biotic observations, mostly on invertebrates (see review in Reed 2002; references in Sedberry 2001; Reed et al. 2006). Studies elsewhere revealed that these deep reefs harbor extensive invertebrate populations composed of hundreds of species (Jensen and Frederiksen 1992; Rogers 1999; Reed 2002). Fish studies related to the deep coral banks are almost non existent (no detailed faunal surveys are published for the western Atlantic), even in the northeastern Atlantic where these corals are better known (Husebo et al. 2002). Although our investigations so far have revealed that many species of fishes and crabs are closely associated with this unique deep-reef habitat, it is unclear whether the deep coral habitat is essential to selected fishes or invertebrates or whether they occupy it opportunistically (see conflicting views in Auster 2005 and Costello et al. 2005). Assessing its 2 significance as fish and invertebrate habitat and addressing the extent to which the deep reef fauna is unique is an important research topic being investigated (S.W. Ross et al., ongoing studies). No deep coral reefs are yet designated as Marine Protected Areas (MPAs) in the US EEZ deeper than 200 m, but if such reefs prove to be important habitat with unique fauna (as they seem to be), they should be candidates for protection as are Oculina coral reefs off east-central Florida. There are a variety of potential threats to the deep coral bottoms: mining, precious coral harvest, disposal activities, petroleum exploration and development, fiber optic and other cable laying, anchoring/gear damage, and fishing activities. MPAs or Habitat Areas of Particular Concern (HAPC) may be viable options for protecting these systems, but considerable data, especially detailed maps, are critical for evaluating how and whether to protect deep coral habitat (Miller 2001). As stated above, publications concerning deep coral banks in the SEUS are limited, particularly for reefs off North Carolina. Much of the data in this review of SEUS deep corals are from ongoing studies of a multi-agency research team (Steve W. Ross, lead Principal Investigator, Univ. North Carolina-Wilmington). Although this research team has collected considerable data on deep coral mounds and has given numerous verbal presentations, publications are still forthcoming. Considering the needs of the South Atlantic Fishery Management Council (SAFMC) to evaluate deep water habitats in a timely manner, the brief descriptions of SEUS deep coral banks (Fig. 1) provided below will serve as an interim tool facilitating potential management options for fragile, productive deep water habitats. This synthesis is preliminary, pending final data analyses and scientific publications, and should be used cautiously with prior consent of the author. Additional information should be obtained from S.W. Ross.
HISTORY OF DEEP CORAL RESEARCH IN THE SEUS
The history of deep coral research in the SEUS is temporally and spatially sporadic. Until recently deep coral research was often a by-product of non coral projects. The major studies that document deep water corals in the area are briefly reviewed. The review below is roughly chronological and not intended to be inclusive. Deep water corals were first reported on the Blake Plateau from 1880 collections of the steamer Blake (Agassiz 1888). These collections were poorly documented, and Agassiz summarized the Blake Plateau bottom as being hard and barren. The research vessel Albatross collected corals on the Blake Plateau in 1886 using beam trawls and tangles. Some of the Lophelia specimens in those collections were deposited in the US National Museum, but were otherwise poorly documented. Much later, Squires (1959) noted several scleractinian species dredged in 1954 off Palm Beach, FL in 686 m. Cairns (1979) corrected coral identifications from Squires (1959) which resulted in the above collection containing Lophelia pertusa, Crispatotrochus (=Caryophyllia) squiresi, Enallopsammia profunda, and Tethocyathus variabilis. An area of very rough topography containing deep corals was described on the Blake Plateau off South Carolina. Many mounds and ridges were surveyed by depth sounder in 1956, 1957, 1959, and 1960 (Stetson et al. 1962). However, thheese features were not confirmed to support extensive
3
A B CF
Figure 1. Ross et al. deep coral study sites (red stars), 2000-2005. CF=Cape Fear.
coral habitat until they were dredged and photographed in 1961 (Stetson 1961). Stetson et al. (1962) gave the first detailed accounting of SEUS coral banks in an area now called the “Stetson Banks” (Fig. 1), confirming that the major hard corals were L. pertusa and Enallopsammia (=Dendrophyllia) profunda. They also reported species of Bathypsammia, Caryophyllia, and Balanophyllia as well as abundant alcyonarians. Additional details from the 1961 cruise, including locations of hundreds of coral mounds, were described by Stetson et al. (1969). Through the 1960s a series of geological papers based largely on precision echosounding data noted that numerous mounds, termed coral mounds, existed on the Blake Plateau and the Florida- Hatteras slope (e.g., Uchupi and Tagg 1966; Uchupi 1967; Zarudzki and Uchupi 1968). Pratt (1968) presented one photograph of Lophelia corals on the Blake Plateau (“Stetson Banks”). In 1967 five manned submersible dives using the DSRV Alvin were made in an area west of the “Stetson Banks”, and two dives confirmed that Enallopsammia (=Dendrophyllia) and Lophelia occurred in certain areas (Milliman et al. 1967). Also from 1967 sampling, Neumann and Ball (1970) described coral topped mounds (to 15 m high) along the slope off Biscayne Bay, FL (around 700-825 m). Although corals were discovered on the Blake Plateau in the 1880s and investigated in the late
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1950s and early 1960s (Squires 1959; Stetson et al. 1962), it seems that such corals were not known off North Carolina until the late 1960s. Based on seismic profiling, Uchupi (1967) first noted the occurrence of a coral mound off Cape Lookout, NC, which may be the same area illustrated (figure caption without further comment) by Rowe and Menzies (1968). Rowe and Menzies (1969) later suggested that Lophelia sp. occurred off the Carolinas in “discontinuous banks” along the 450 m contour, but gave no specific data. Likewise, Menzies et al. (1973) vaguely referenced a “Lophohelia” bank off Cape Lookout, repeating a figure in Rowe and Menzies (1969) and presenting a bottom photograph of a reef in 458 m. Cairns (1979) plotted a locality off Cape Lookout in his distribution map for Lophelia without comment. Aside from Uchupi (1967), the above North Carolina records seem to have originated from a training cruise of the R/V Eastward (E-25-66, I.E. Gray, chief scientist) during which a coral bank was photographed by drop camera (station E-4937, 475 m) and dredged (E-4933, 425 m) on 30 June 1966. The Menzies et al. (1973, Fig. 4-4 B) photograph is from that cruise. This coral bank was discovered accidentally (independently of Uchupi 1967) as a result of constantly running the R/V Eastward’s depth sounder (L. McCloskey and G. Rowe, pers. comm.). There were a few other short Eastward cruises to this area off Cape Lookout under direction of Menzies, Rowe, Gray, or McCloskey but no coral data were published. This Eastward station area was trawled and surveyed by sonar in May 1983 (R/V Delaware II cruise, S.W. Ross, chief scientist), but no hard bottom or coral were found. Coral mounds were located in this vicinity during an undersea survey using the Navy’s NR-1 nuclear research submersible (15-18 Nov 1993, K.J. Sulak and S.W. Ross, unpubl. data). To date three major coral mounds have been located and studied off North Carolina (Reed and Ross 2005; S.W. Ross et al., unpubl. data), and several other mounds may exist. The slope off Cape Lookout appears to be the northern extent of deep sea, cold water corals in the SEUS region. References for the SEUS deep coral areas continued to result from studies that were generally not directed toward corals or that were geological in nature. Exceptions include Cairns (1979, 1981, 2000, 2001a), who listed ranges for a number of deep sea Scleractinia and azooxanthellate corals in this area, relying mostly on museum records. From five Alvin dives in 1971 in the eastern Florida Straits off Little Bahama Bank, Neumann et al. (1977) described hard carbonate mounds that were covered in various corals (Lophelia and Enallopsammia) and other invertebrates, and coined the term “lithoherms” for these structures. In this same area in 1982 and also using Alvin, several coral species were collected and aged, indicating that these animals lived from several hundred up to 1800 years (Griffin and Druffel 1989; Druffel et al. 1990, 1995). Since these corals have annual rings that contain a wealth of information about past climates, ocean productivity, and contamination, this significant discovery has vast implications for the scientific value of deep sea corals. During a study of surficial and deeper sediments of the Florida-Hatteras slope and inner Blake Plateau, Ayers and Pilkey (1981) documented a number of coral banks, collected corals, and dated several coral samples. Depending on location in a core, their dead coral samples ranged in age from 5,000 to 44,000 years old. They dated a living specimen at 680 years old, but suggested that this age probably reflected age of the carbon pool in the surrounding water. Pinet et al. (1981) also mapped coral banks overlapping the same area as Ayers and Pilkey (1981). Blake et al. (1987) briefly mentioned some soft and hard coral occurrences on the Blake Plateau. Many deep reef locations were suggested by the U.S. Geological Survey sidescan sonar mapping (cruises in 1987) of the continental slope (EEZ-SCAN 87 Scientific Staff 1991); however, this large scale geological survey had little habitat verification. Perhaps the first study to document the invertebrate community associated with deep coral habitat in this area reviewed biozonation of lithoherms in the northeastern Straits of Florida (Messing et al. 1990). Genin et al. (1992) noted that sponges and gorgonians were common along the outer Blake escarpment (2624-4016 m) based on 1980 Alvin dives. They suggested that these communities were unusually dense for sites lacking sediment.
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Popenoe (1994) discussed the distribution and formation of coral mounds on the Blake Plateau and presented a few bottom photographs. Paull et al. (2000) surveyed deep coral habitats off the Florida-Georgia border, dated parts of the structures, and suggested that such habitat was very common. Their dating indicated that some mounds may range from 18,000 to 33,000 years old. Popenoe and Manheim (2001) extensively reviewed geology, history, and habitats of a portion of the Blake Plateau around the Charleston Bump, discussing various parameters that may control coral mound formation. Wenner and Barans (2001) described benthic habitats of the Charleston Bump area and noted some of the invertebrates and fishes occurring with deep corals. George (2002) also discussed a coral habitat southeast of Cape Fear, NC (“Agassiz Coral Hills”) in 650-750 m dominated by Bathypsammia tintinnabulum. Apparently the B. tintinnabulum used by Emilini et al. (1978) came from the area and collections described by George (2002). Reed (2002a, b, 2006) described several large areas of deep corals on the Blake Plateau. As part of a SEAMAP bottom mapping project, data to be scanned for evidence of deep corals in this area were summarized by Arendt et al. (2003). Beginning in 2000 and continuing through the present, deep coral (or related habitat) research in the SEUS was stimulated by funding of several studies through the NOAA Office of Ocean Exploration (supplemented by other sources). Teams lead by Principal Investigators S. Brooke, S. Pomponi, S.W. Ross, and G.R. Sedberry explored deep coral banks throughout the SEUS, mapping habitats, cataloging fauna, and conducting basic biological studies. A multi-investigator effort to create detailed habitat classifications (Southeastern US Deep-Sea Corals initiative, SEADESC) from past submersible dives in the area is underway. A related effort to generally locate hard bottom or coral habitat between 200 and 2000 m (SEAMAP) is also underway. Future publications should be forthcoming from the considerable data collected by these efforts.
DEEP SEA CORALS OF THE SEUS
The SEUS slope area, including the slope off the Florida Keys, appears to have a unique assemblage of deep water Scleractinia (Cairns and Chapman 2001). The warm temperate assemblage identified by Cairns and Chapman (2001) contained about 62 species, four endemic to the region. This group was characterized by many free living species, few species living deeper than 1000 m, and many species with amphi-Atlantic distributions. Based on literature the SEUS region contains at least 109 species of deep corals (classes Hydrozoa and Anthozoa, Ross and Nizinski in press). This number is conservative, since collection of corals has rarely been a research priority. Lophelia pertusa, the major structure building coral in the deep sea, is fragile and susceptible to physical destruction (Fossa et al. 2002). Lophelia reefs are widespread, occurring not only on the SEUS slope, but also in the Gulf of Mexico, off Nova Scotia, in the northeastern Atlantic, the South Atlantic, the Mediterranean, Indian Ocean and in parts of the Pacific Ocean over a depth range of 50 to 2170 m (Cairns 1979; Rogers 1999). Coral habitats dominated by Lophelia pertusa are common throughout the SEUS in depths of about 370 to at least 800 m. Reed and Ross (2005) summarized area deep coral research. While their study areas do not cover all known deep coral habitat in the region, they have conducted work over most of the well known coral sites (Fig. 1). Although Lophelia may occur in small scattered colonies attached to various hard substrates, it also forms complex, high profile features (bioherms). The ridges and reef mounds, some rising more than 100 m from the open substrate, accelerate bottom currents which are favorable to attached filter-feeders. Thus, the growing reef alters local currents, enhancing the environment for continued coral growth and faunal recruitment (Genin et al. 1986). Along the sides and around the bases of these banks are rubble zones of dead coral pieces which may extend large distances away from the mounds.
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Data are lacking on how Lophelia coral banks form despite several hypotheses (Hovland et al. 1998; Hovland and Risk 2003; Masson et al. 2003). The mounds off North Carolina and those in some other SEUS locations (particularly East of South-central Florida) appear to be formed by successive coral growth, collapse, and sediment entrapment (Wilson 1979; Ayers and Pilkey 1981; Paull et al. 2000; Popenoe and Manheim 2001). Other deep coral habitats in the area (especially on the Blake Plateau) seem to be formed by coral colonization of appropriate hard substrates, without mound formation by the corals. Bottom currents that are too strong may prevent mound formation (Popenoe and Manheim 2001) because sediments cannot be trapped. Assuming currents also carry appropriate foods, it may be that currents with variable speeds or at least currents of moderate speeds (fast enough to facilitate filter feeding but not too fast to prevent sediment entrapment) coupled with a supply of sediment are the conditions necessary to facilitate coral mound formation (Rogers 1999). Regardless of how formed, elevated topography appears to be an important attribute for well developed coral communities (Masson et al. 2003). Although exactly how these corals feed and grow are poorly known, data indicate that food sources are not chemosynthetic and are probably surface derived (Duineveld et al. 2004). These deep reefs may be hundreds to tens of thousands of years old (Neumann et al. 1977; Wilson 1979; Ayers and Pilkey 1981; Mikkelsen et al. 1982; Mortensen and Rapp 1998); however, aging data are so limited (especially in the western Atlantic) that the distribution of coral mound ages in the western Atlantic is unclear. Regardless, it seems likely that most of these structures are at least thousands of years old. While the genetic structure (gene flow, population relationships, taxonomic relationships) of Lophelia in the northeastern Atlantic has been described (Le Goff-Vitry et al. 2004), such studies are just beginning in the western Atlantic ©. Morrison et al., unpubl. data). Bamboo (Family Isididae, four species) and black corals (Families Leiopathidae and Schizopathidae, ca. four species) are also important structure forming corals in the SEUS (Fig. 2). These corals occur locally in moderate abundances, but their distributions seem to be limited to the region south of Cape Fear, NC. Colonies may reach heights of 1-2 m. Bamboo and black coral colonies, occurring either singly or in small aggregations, may be observed either in association with hard coral colonies or as separate entities. Furthermore, some of these living components of the deep reefs (e.g., black corals, zoanthids) are hundreds to thousands of years old (Griffin and Druffel 1989; Druffel et al. 1995; Williams et al. in press; C. Holmes and S.W. Ross, unpubl. data), the oldest animals on Earth. They form annual or regular bands and these bands contain important chemical records on past climates, ocean physics, ocean productivity, pollution, and data relevant to global geochemical cycles. A major effort to investigate these geochemical data is being started by USGS ©. Holmes and S.W. Ross)
NORTH CAROLINA DEEP CORAL BANKS
Off North Carolina, Lophelia forms what may be considered classic mounds (three areas surveyed so far) that appear to be a sediment/coral rubble matrix topped with almost monotypic stands of L. pertusa (Figs. 3-4). Although Lophelia is the dominant hard coral off North Carolina, other scleractinians contribute to the overall complexity of the habitat. These include the colonial
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Figure 2. Selected views of Black corals and Bamboo corals on the Blake Plateau (Ross et al., unpubl. data).
8 corals Madrepora oculata and Enallopsammia spp. as well as a variety of solitary corals. These hard corals tend to live on or within the Lophelia matrix. The three North Carolina Lophelia mounds are the northernmost coral banks in the SEUS. Because these banks seem to be a northern terminus for a significant zoogeographic region, they may be unique in biotic resources as well as habitat expression. The three NC banks are generally similar in physical attributes and faunal composition. Some observed differences, however, are being investigated, and more detailed results will be presented in several peer reviewed publications in preparation (Ross et al.). For convenience these three areas have been designated as Cape Lookout Lophelia Bank A, Cape Lookout Lophelia Bank B, and Cape Fear Lophelia Bank. These names are to facilitate research and may eventually be changed. General descriptions of the NC coral mounds and associated fauna follows. Since there is almost no data published for the NC deep coral banks and because they are different than those to the south, they are discussed in more detail below. Several potential deep coral banks were identified in the USGS survey of the EEZ off of North Carolina (EEZ-SCAN 87 Scientific Staff 1991). Attempts were made (Ross et al. cruises) to locate a few of these banks to no avail. These coral mounds, especially off southern North Carolina, would be important to document as they would occur in what may be a transition area between a region of coral/sediment built mounds composed almost entirely of L. pertusa and the area to the south where coral development is generally quite different.
Biological Communities of the North Carolina Coral Banks
Fish communities are extensive but difficult to document on and around these coral banks. Some level of commercial fishing activity seems to occur on the NC Banks, as we have observed trash and entangled fishing gear on the reefs. Because the fish data have been extensively analyzed and are nearly ready for submission for publication, a more detailed treatment of the region’s fish data is presented below. An impressive biological aspect of these coral mounds (aside from the corals themselves) is the well developed and abundant invertebrate fauna. We have not yet detected major differences in the invertebrate fauna among the three North Carolina banks. Galatheid crabs (especially Eumunida picta) and the brisingid basket star (Novodinia antillensis) were particularly obvious, perching high in coral bushes to catch passing animals or filter in the currents (Fig. 5). One very different aspect of the North Carolina deep coral habitat compared to the rest of the South Atlantic Bight is the massive numbers of a brittle star (Ophiacantha bidentata) covering both dead and living coral colonies. They are perhaps the most abundant macroinvertebrate on these banks and may constitute a major food source (Brooks et al. in review). In places the bottom is covered with huge numbers of several species of anemones (Fig. 5). The abundance of filter feeders suggest a food rich habitat.
Cape Lookout Lophelia Bank A
Aside from a few maps (see above) there are no published data from this coral mound. This area was apparently first occupied by the R/V Eastward (see above) which gave a location of 34 18' N, 75 48' W. Two trawl stations and a sonar survey of the Eastward station area in May 1983 using the R/V Delaware II (S.W. Ross, chief scientist), revealed no indication of hard bottom or coral. The Eastward navigated with LORAN A, and since their station was about 1-1.5 nmi (2-2.7
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Figure 3. Selected views of Lophelia pertusa habitat and depth sounder recordings for the two deep coral mounds off Cape Lookout, NC (Ross et al., unpubl. data).
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Figure 4. Selected views of the deep coral mound off Cape Fear, NC. Bottom panel is a 3-D reconstruction of this feature with general habitat classifications (SEADESC) from JSL dives (S.W. Ross, unpubl. data).
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Figure 5. Various invertebrates common on SEUS deep coral banks. From left to right and top to bottom: Eumunida picta perched on Lophelia pertusa, Rochina crassa sitting on dead Lophelia, close up of Echinus urchin with brittle stars (Ophiacantha bidentata) among coral branches, anemone Actinauge, basket star Novodinia antillensis, two anemonies and brittle stars among coral branches. These photographs were from North Carolina coral banks (S.W. Ross et al. unpubl. data).
12 km) from the large coral bank area sampled later (Fig. 3), it is likely that the less accurate LORAN may have put the Eastward station off of the actual reef. However, the possibility that a reef does exist on or near the E-4937 station cannot be discounted without a more detailed survey of that location. The USGS side scan survey (EEZ-SCAN 87 Scientific Staff 1991) illustrated reefs in this area, and coordinates from that survey guided a cruise using the Navy’s NR-1 nuclear research submersible (Sulak and Ross, unpubl. data) during 15-18 Nov 1993. However, there also seemed to be a navigation issue with this cruise in that locations plotted from the NR-1 track are offset about 0.5 nmi (1 km) from the large mounds later located by Ross et al. (unpubl. data). A later ship sonar survey of the NR-1 locations did not yield obvious reef areas. Between summer 2000 and fall 2005 Ross et al. (unpubl. data) sampled this area extensively using a variety of methods throughout the water column. Their major method for collecting bottom data on the reef proper was the JSL research submersible. Seventeen dives were made on coral mounds in this area (Fig. 6, Table 1), and observations from these totaled nearly 37 hours (bottom time). Preliminary observations suggest that this area contains the most extensive coral mounds off North Carolina; however, data are lacking to adequately judge overall sizes and areal coverage. Ross et al. JSL dives in this area ranged from 370-447 m (Table 1). Mean bottom temperatures ranged from 6.3 to 10.9 C, while mean bottom salinities were always around 35 (Table 2). There appear to be several prominences capping a ridge system, thus, presenting a very rugged and diverse bathymetry (Fig. 6), but there are also other mounds away from the main ridge sampled (Fig. 6). The main mound system rises vertically nearly 80 m over a distance of about 1 km, and in places exhibits slopes in excess of 50-60 degrees. Sides and tops of these mounds are covered with extensive colonies of living Lophelia pertusa (Fig. 3), with few other corals being observed. Dead colonies and coral rubble interspersed with sandy channels are also abundant. Extensive coral rubble zones surround the mounds for a large, but unknown, distance (exact area not yet surveyed), especially at the bases of the mounds/ridges, and in places seem to be quite thick. These mounds appear to be formed by successive coral growth, collapse, and sediment entrapment (Wilson 1979; Popenoe and Manheim 2001). These topographic highs accelerate bottom currents which favor attached filter-feeders, and very strong bottom currents have been observed.
Cape Lookout Lophelia Bank B
Except for a few maps (see above) there are no published data from this coral mound. The USGS side scan survey (EEZ-SCAN 87 Scientific Staff 1991) illustrated reefs in this area, and, as above, coordinates from that survey guided the cruise using the NR-1 nuclear research submersible (Sulak and Ross, unpubl. data) during 15-18 Nov 1993. The same navigation issue with this cruise described above was also apparent in this area, NR-1 stations being about 1-1.4 nmi (2-2.6 km) from major mounds located later by Ross et al. Between summer 2001and fall 2005 Ross et al. (unpubl. data) sampled this area using a variety of methods throughout the water column. The JSL submersible was the major method for collecting bottom data on the reef proper. Nine dives were made on coral mounds in this area (Fig. 7, Table 1), and observations from these totaled about 20 hours. The least amount of data are available for this area. Mounds appear to cover a smaller area than those described above, but here again better mapping data are needed. Ross et al. JSL dives in this area ranged from 375-450 m (Table 1). Mean bottom temperatures ranged from 5.8 to 10.4 C, and as above mean bottom salinities were always around 35 (Table 2). These mounds rise at least
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Table 1. Johnson-Sea-Link (JSL) research dives conducted on deep coral habitat on the slope of the southeastern US by S.W. Ross et al., summer 2000-fall 2005. Start, end and total times represent bottom times in minutes. Station Date Location Start End Total Start Start End End Start-End Time Time Time (min) Lat Long Lat Long Depth (m)
JSLI-2000-4206 28-Jul-00 Cape Lookout A 08:42 10:36 114 34° 19.52 75° 47.05 34° 19.45 75° 47.25 430-389 JSLI-2000-4207 28-Jul-00 Cape Lookout A 15:56 17:45 109 34° 19.57 75° 47.13 34° 19.42 75° 47.29 418-405 JSLI-2001-4361 22-Sep-01 Cape Lookout A 08:44 11:23 159 34° 19.68 75° 47.37 34° 19.69 75° 47.53 427-384 JSLI-2001-4362 22-Sep-01 Cape Lookout A 16:21 18:36 135 34° 19.43 75° 47.49 34° 19.42 75° 47.51 399-370 JSLI-2001-4363 23-Sep-01 Cape Lookout A 09:02 11:15 133 34° 19.42 75° 47.45 34° 19.41 75° 47.50 417-371 JSLI-2001-4364 23-Sep-01 Cape Lookout A 16:02 18:53 171 34° 18.84 75° 47.01 34° 18.77 75° 47.13 442-398 JSLI-2001-4365 24-Sep-01 Cape Lookout B 08:42 11:15 153 34° 11.34 75° 53.80 34° 11.41 75° 53.74 431-414 JSLI-2001-4366 24-Sep-01 Cape Lookout B 16:18 17:32 74 34° 10.75 75° 53.51 34° 10.77 75° 53.37 449-437 JSLII-2002-3304 11-Aug-02 Cape Lookout A 08:33 11:01 148 34° 19.71 75° 47.04 34° 19.51 75° 46.21 447-386 JSLII-2002-3305 11-Aug-02 Cape Lookout A 16:30 18:59 149 34° 19.46 75° 47.20 34° 19.48 75° 47.20 416-385 JSLII-2002-3306 12-Aug-02 Cape Lookout A 08:32 10:59 147 34° 19.4 75° 47.2 34° 19.45 75° 47.25 418-384 JSLII-2002-3307 12-Aug-02 Cape Lookout A 16:24 17:11 47 34° 19.48 75° 47.45 34° 19.50 75° 47.55 416-383 JSLII-2002-3308 13-Aug-02 Cape Fear 08:29 10:58 149 33° 34.33 76° 29.05 33° 34.43 76° 27.90 449-373 JSLII-2003-3419 17-Aug-03 Stetson 08:40 10:51 131 32° 01.75 77° 40.44 32° 02.01 77° 40.49 622-597 JSLII-2003-3420 17-Aug-03 Stetson 16:18 18:24 126 32° 02.01 77° 40.71 32° 02.04 77° 40.93 626-629 JSLII-2003-3425 21-Aug-03 Cape Fear 08:21 10:47 146 33° 34.38 76° 27.93 33° 34.46 76° 27.87 386-379 JSLII-2003-3426 21-Aug-03 Cape Fear 16:36 19:03 147 33° 34.38 76° 27.91 33° 34.33 76° 27.91 371-377 JSLII-2003-3427 22-Aug-03 Cape Fear 08:33 10:51 138 33° 34.28 76° 27.75 33° 34.48 76° 27.70 381-418 JSLII-2003-3428 22-Aug-03 Cape Fear 16:11 18:17 126 33° 34.38 76° 27.95 33° 34.44 76° 27.89 377-371 JSLII-2003-3429 23-Aug-03 Cape Lookout B 08:54 11:10 136 34° 11.15 75° 54.03 34° 11.42 75° 53.75 435-415 JSLII-2003-3430 23-Aug-03 Cape Lookout A 16:24 18:59 155 34° 19.37 75° 47.33 34° 19.40 75° 47.25 415-394 JSLII-2003-3431 24-Aug-03 Cape Lookout A 08:36 10:52 136 34° 19.52 75° 47.04 34° 19.42 75° 47.24 432-389 JSLII-2003-3432 24-Aug-03 Cape Lookout A 16:47 18:57 130 34° 19.43 75° 47.16 34° 19.48 75° 47.21 424-385 JSLI-2004-4681 09-Jun-04 North Cape Canaveral 09:10 11:12 122 28º 47.55 79º 37.19 28º 47.60 79º 37.31 783-709 JSLI-2004-4682 09-Jun-04 North Cape Canaveral 17:06 19:08 122 28º 47.76 79º 37.30 28º 47.75 79º 37.24 770-760 JSLI-2004-4683 10-Jun-04 Jacksonville Lithoherms 08:32 10:55 143 30º 31.05 79º 39.62 30º 30.97 79º 39.72 568-544 JSLI-2004-4684 10-Jun-04 Jacksonville Lithoherms 16:37 18:43 126 30º 30.94 79º 39.62 30º 30.84 79º 39.62 569-554 JSLI-2004-4685 11-Jun-04 Jacksonville Lithoherms 08:45 11:00 135 30º 48.81 79º 37.81 30º 48.70 79º 37.93 652-636 JSLI-2004-4686 11-Jun-04 Jacksonville Lithoherms 17:02 18:55 113 30º 30.13 79º 39.09 30º 30.10 79º 39.18 638-593 JSLI-2004-4687 12-Jun-04 Savannah Banks 08:32 10:13 101 31º 44.36 79º 06.09 31º 44.52 79º 05.66 540-497 JSLI-2004-4688 12-Jun-04 Savannah Banks 16:27 18:00 93 31º 46.45 79º 11.70 31º 46.56 79º 11.59 532-516 JSLI-2004-4689 13-Jun-04 Stetson 08:37 10:37 120 31º 49.15 77º 36.77 31º 49.15 77º 36.20 672-668 JSLI-2004-4692 15-Jun-04 Cape Lookout A 08:29 10:33 124 34º 19.43 75º 47.17 34º 19.44 75º 47.22 425-384 JSLI-2004-4693 15-Jun-04 Cape Lookout A 16:20 18:27 127 34º 19.44 75º 47.14 34º 19.51 75º 47.15 431-392 JSLI-2004-4694 16-Jun-04 Cape Lookout B 08:29 10:41 132 34º 11.28 75º 53.62 34º 11.28 75º 53.79 440-396 JSLI-2004-4695 16-Jun-04 Cape Lookout B 16:49 18:59 130 34º 11.41 75º 53.65 34º 11.41 75º 53.74 442-414
14
JSLI-2004-4696 17-Jun-04 Cape Fear 08:31 10:25 114 33º 34.37 76º 27.71 33º 34.36 76º 27.67 390-402 JSLI-2004-4697 17-Jun-04 Cape Fear 16:42 18:24 102 33º 34.57 76º 27.83 33º 34.59 76º 27.77 405-411 JSLI-2004-4698 18-Jun-04 Stetson 09:42 11:31 109 31º 49.45 77º 36.69 31º 49.56 77º 36.79 703-664 JSLI-2004-4699 18-Jun-04 Stetson 16:59 19:09 130 31º 50.89 77º 36.72 31º 50.75 77º 36.77 696-660 JSLI-2004-4700 19-Jun-04 Jacksonville Lithoherms 09:37 11:07 90 30º 30.76 79º 39.68 30º 30.85 79º 39.60 564-558 JSLI-2004-4701 19-Jun-04 Jacksonville Lithoherms 17:04 18:43 99 30º 28.94 79º 38.50 30º 28.93 79º 38.38 647-674 JSLI-2004-4702 20-Jun-04 North Cape Canaveral 08:38 10:42 124 28º 47.70 79º 37.40 28º 47.61 79º 37.38 738-713 JSLI-2004-4703 20-Jun-04 North Cape Canaveral 17:08 18:52 104 28º 46.62 79º 36.96 28º 46.62 79º 36.96 756-742 JSLI-2004-4704 21-Jun-04 South Cape Canaveral 08:37 10:41 124 28º 02.64 79º 36.82 28º 02.53 79º 36.75 739-738 JSLI-2004-4705 21-Jun-04 South Cape Canaveral 17:18 19:08 110 28º 02.16 79º 36.84 28º 02.38 79º 36.78 725-689 JSLI-2005-4890 17-Oct-05 Cape Lookout A 08:36 10:43 127 34º 19.59 75º 47.09 34º 19.47 75º 47.22 420-389 JSLI-2005-4891 17-Oct-05 Cape Lookout A 16:32 18:27 115 34º 19.49 75º 47.44 34º 19.37 75º 47.56 433-380 JSLI-2005-4892 18-Oct-05 Cape Lookout B 08:22 10:42 140 34º 13.90 75º 52.44 34º 14.08 75º 52.33 411-375 JSLI-2005-4893 18-Oct-05 Cape Lookout B 16:30 18:31 121 34º 14.00 75º 52.30 34º 14.19 75º 52.28 418-371 JSLI-2005-4894 19-Oct-05 Cape Lookout B 08:22 10:59 157 34º 10.66 75º 53.59 34º 11.00 75º 53.36 450-409 JSLI-2005-4895 19-Oct-05 Cape Lookout B 16:22 18:51 149 34º 12.96 75º 53.09 34º 12.96 75º 53.02 413-395 JSLI-2005-4896 20-Oct-05 Cape Fear 08:24 10:50 146 33º 34.18 76º 27.89 33º 34.17 76º 27.77 397-375 JSLI-2005-4897 20-Oct-05 Cape Fear 16:21 18:26 125 33º 34.64 76º 27.98 33º 34.65 76º 27.95 443-408 JSLI-2005-4898 21-Oct-05 Stetson North 08:37 10:35 118 32º 15.94 77º 28.42 32º 16.17 77º 28.47 642-550 JSLI-2005-4899 21-Oct-05 Stetson North 16:22 18:25 123 32º 15.84 77º 28.82 32º 15.83 77º 29.02 603-587 JSLI-2005-4900 22-Oct-05 Savannah Banks 17:03 19:17 134 31º 44.36 79º 06.16 31º 44.57 79º 05.53 543-519 JSLI-2005-4901 23-Oct-05 Savannah Banks 08:28 08:35 7 31º 42.36 79º 07.42 31º 42.30 79º 07.39 508-507 JSLI-2005-4902 26-Oct-05 Savannah Banks 16:39 18:58 139 31º 42.26 79º 07.88 31º 42.32 79º 07.31 516-514 JSLI-2005-4903 27-Oct-05 Stetson 08:30 10:21 111 32º 01.12 77º 40.00 32º 00.95 77º 40.16 633-633 JSLI-2005-4904 27-Oct-05 Stetson 16:29 18:52 143 31º 50.81 77º 36.83 31º 50.79 77º 36.74 652-657 JSLI-2005-4905 30-Oct-05 Savannah Banks 16:12 18:35 143 31º 46.91 79º 12.26 31º 46.43 79º 12.10 541-515 JSLI-2005-4906 30-Oct-05 Savannah Banks 17:26 18:52 86 31º 46.49 79º 11.64 31º 46.62 79º 11.56 525-515 JSLI-2005-4907 01-Nov-05 Jacksonville Lithoherms 08:28 10:47 139 30º 48.15 79º 38.39 30º 48.03 79º 38.50 534-530 JSLI-2005-4908 01-Nov-05 Jacksonville Lithoherms 16:38 18:55 137 30º 31.12 79º 39.63 30º 31.26 79º 39.41 585-625
15
Table 2. Bottom temperature and salinity data from Johnson-Sea-Link (JSL) dives on deep coral habitat on the slope of the southeastern US (S.W. Ross et al. unpubl. data).
Station Date Location Mean Temp Temp Range Mean Salinity (C°) ± SE (C°) Salinit y ± SE Range
JSLI-2000-4206 28-Jul-00 Cape Lookout A 8.49 ± 0.02 5.64-10.64 35.20 ± 0.00 34.04-36.20 JSLI-2000-4207 28-Jul-00 Cape Lookout A 8.63 ± 0.01 6.23-9.44 35.20 ± 0.00 34.06-35.81 JSLI-2001-4361 22-Sep-01 Cape Lookout A 9.49 ± 0.00 9.09-9.92 35.22 ± 0.00 35.02-35.60 JSLI-2001-4362 22-Sep-01 Cape Lookout A 10.13 ± 0.00 9.22-10.57 35.31 ± 0.00 34.99-35.70 JSLI-2001-4363 23-Sep-01 Cape Lookout A 10.44 ± 0.00 9.90-10.80 35.35 ± 0.00 35.11-35.52 JSLI-2001-4364 23-Sep-01 Cape Lookout A 10.06 ± 0.01 9.00-10.86 35.30 ± 0.00 35.03-35.53 JSLII-2002-3304 11-Aug-02 Cape Lookout A 9.61 ± 0.01 6.30-10.88 35.26 ± 0.00 33.91-36.03 JSLII-2002-3305 11-Aug-02 Cape Lookout A 9.24 ± 0.00 8.97-10.12 35.21 ± 0.00 34.70-35.69 JSLII-2002-3306 12-Aug-02 Cape Lookout A 10.90 ± 0.01 8.87-14.85 35.39 ± 0.00 34.02-36.09 JSLII-2002-3307 12-Aug-02 Cape Lookout A 10.15 ± 0.00 9.83-10.54 35.30 ± 0.00 34.99-35.49 JSLII-2003-3430 23-Aug-03 Cape Lookout A 6.33 ± 0.00 5.90-6.88 35.06 ± 0.00 34.90-35.56 JSLII-2003-3431 24-Aug-03 Cape Lookout A 7.08 ± 0.01 6.20-8.29 35.08 ± 0.00 34.92-35.28 JSLII-2003-3432 24-Aug-03 Cape Lookout A 8.27 ± 0.00 7.45-9.04 35.13 ± 0.00 34.81-35.31 JSLI-2004-4692 15-Jun-04 Cape Lookout A 9.81 ± 0.00 9.55-9.99 35.28 ± 0.00 35.19-35.36 JSLI-2004-4693 15-Jun-04 Cape Lookout A 9.11 ± 0.00 8.04-9.57 35.20 ± 0.00 35.02-35.34 JSLI-2005-4890 17-Oct-05 Cape Lookout A 8.14 ± 0.01 5.51-8.98 35.13 ±0.00 34.89-35.32 JSLI-2005-4891 17-Oct-05 Cape Lookout A 9.03 ± 0.00 8.36-9.6 35.19 ± 0.00 35.06-35.36
JSLI-2001-4365 24-Sep-01 Cape Lookout B 10.01 ± 0.00 9.58-10.30 35.27 ± 0.00 35.13-35.41 JSLI-2001-4366 24-Sep-01 Cape Lookout B 9.81 ± 0.00 9.61-10.14 35.25 ± 0.00 35.11-35.43 JSLII-2003-3429 23-Aug-03 Cape Lookout B 5.82 ± 0.00 5.42-5.97 35.04 ± 0.00 34.99-35.12 JSLI-2004-4694 16-Jun-04 Cape Lookout B 10.43 ± 0.01 9.39-11.19 35.36 ± 0.00 35.20-35.53 JSLI-2004-4695 16-Jun-04 Cape Lookout B 9.95 ± 0.00 9.70-11.34 35.32 ± 0.00 35.02-35.83 JSLI-2005-4892 18-Oct-05 Cape Lookout B 8.77 ± 0.00 8.64-9.73 35.13 ± 0.00 35.01-35.32 JSLI-2005-4893 18-Oct-05 Cape Lookout B 9.12 ± 0.00 8.42-9.60 35.16 ± 0.00 35.04-35.30 JSLI-2005-4894 19-Oct-05 Cape Lookout B 7.55 ± 0.00 6.30-8.24 35.07 ± 0.00 34.96-35.22 JSLI-2005-4895 19-Oct-05 Cape Lookout B 7.77 ± 0.00 7.63-7.93 35.04 ± 0.00 34.98-35.04
JSLII-2002-3308 13-Aug-02 Cape Fear 9.13 ± 0.00 8.42-9.53 35.18 ± 0.00 34.80-35.45 JSLII-2003-3425 21-Aug-03 Cape Fear 9.54 ± 0.00 9.54-9.72 35.20 ± 0.00 35.10-35.34 JSLII-2003-3426 21-Aug-03 Cape Fear 10.18 ± 0.01 9.25-11.22 35.29 ± 0.00 35.00-35.60 JSLII-2003-3427 22-Aug-03 Cape Fear 8.69 ± 0.00 7.93-9.83 35.15 ± 0.00 34.75-35.61 JSLII-2003-3428 22-Aug-03 Cape Fear 9.13 ± 0.00 8.68-9.70 35.19 ± 0.00 35.14-35.26 JSLI-2004-4696 17-Jun-04 Cape Fear 9.10 ± 0.00 9.00-9.54 35.14 ± 0.00 35.05-35.30 JSLI-2004-4697 17-Jun-04 Cape Fear 11.70 ± 0.00 11.01-12.09 35.48 ± 0.00 35.33-35.67 JSLI-2005-4896 20-Oct-05 Cape Fear 8.06 ± 0.00 7.91-8.26 35.06 ± 0.00 35.01-35.10 JSLI-2005-4897 20-Oct-05 Cape Fear 8.00 ± 0.00 7.78-8.23 35.06 ± 0.00 34.98-35.12
JSLI-2005-4898 21-Oct-05 Stetson North 7.97 ± 0.00 7.22-8.10 35.09 ± 0.00 35.03-35.17 JSLI-2005-4899 21-Oct-05 Stetson North 8.64 ± 0.00 8.13-9.91 35.14 ± 0.00 35.01-35.32
JSLII-2003-3419 17-Aug-03 Stetson 10.89 ± 0.00 10.78-11.03 35.39 ± 0.00 35.37-35.41 JSLII-2003-3420 17-Aug-03 Stetson 9.91 ± 0.00 9.83-10.06 35.25 ± 0.00 35.23-35.27 JSLI-2004-4689 13-Jun-04 Stetson 12.20 ± 0.00 12.12-12.30 35.55 ± 0.00 35.51-35.60 JSLI-2004-4698 18-Jun-04 Stetson 11.00 ± 0.00 10.94-11.82 35.36 ± 0.00 35.31-35.54 JSLI-2004-4699 18-Jun-04 Stetson 10.97 ± 0.00 10.93-11.13 35.36 ± 0.00 35.27-35.47 JSLI-2005-4903 27-Oct-05 Stetson 7.57 ± 0.00 7.34-8.05 35.18 ± 0.00 35.14-35.22 JSLI-2005-4904 27-Oct-05 Stetson 9.71 ± 0.00 8.19-11.76 35.30 ± 0.00 34.92-35.79
16
JSLI-2004-4687 12-Jun-04 Savannah Banks 9.07 ± 0.00 8.97-9.13 35.12 ± 0.00 35.09-35.14 JSLI-2004-4688 12-Jun-04 Savannah Banks 8.20 ± 0.00 8.18-8.26 35.02 ± 0.00 35.00-35.04 JSLI-2005-4900 22-Oct-05 Savannah Banks 9.18 ± 0.00 9.06-9.26 35.16 ± 0.00 35.02-35.18 JSLI-2005-4901 23-Oct-05 Savannah Banks … … … … JSLI-2005-4902 26-Oct-05 Savannah Banks 8.11 ± 0.00 8.08-8.21 35.02 ± 0.00 35.00-35.05 JSLI-2005-4905 30-Oct-05 Savannah Banks 7.69 ± 0.00 7.53-7.84 35.01 ± 0.00 34.97-35.04 JSLI-2005-4906 30-Oct-05 Savannah Banks 7.37 ± 0.00 7.36-7.56 34.96-35.04 35.00 ± 0.00
JSLI-2004-4683 10-Jun-04 Jacksonville 10.53 ± 0.00 10.34-10.90 35.28 ± 0.00 35.13-35.48 Lithoherms JSLI-2004-4684 10-Jun-04 Jacksonville 9.63 ± 0.01 9.04-10.50 35.18 ± 0.00 34.85-35.50 Lithoherms JSLI-2004-4685 11-Jun-04 Jacksonville 7.84 ± 0.00 7.80-7.98 34.99 ± 0.00 34.93-35.03 Lithoherms JSLI-2004-4686 11-Jun-04 Jacksonville 9.91 ± 0.00 9.80-10.02 35.21 ± 0.00 35.17-35.26 Lithoherms JSLI-2004-4700 19-Jun-04 Jacksonville 7.64 ± 0.00 7.52-8.34 34.97 ± 0.00 34.88-35.08 Lithoherms JSLI-2004-4701 19-Jun-04 Jacksonville 7.37 ± 0.00 7.31-7.50 34.95 ± 0.00 34.92-34.99 Lithoherms JSLI-2005-4907 01-Nov-05 Jacksonville 7.91 ± 0.00 7.46-8.29 35.02 ± 0.00 34.94-35.09 Lithoherms JSLI-2005-4908 01-Nov-05 Jacksonville 7.33 ± 0.00 7.22-7.61 34.98 ± 0.00 34.94-35.00 Lithoherms
JSLI-2004-4681 09-Jun-04 North Cape 6.75 ± 0.00 6.73-6.89 34.90 ± 0.00 34.86-34.94 Canaveral JSLI-2004-4682 09-Jun-04 North Cape 6.80 ± 0.00 6.78-6.96 34.90± 0.00 34.84-34.99 Canaveral JSLI-2004-4702 20-Jun-04 North Cape 6.55 ± 0.00 6.54-6.65 34.91 ± 0.00 34.87-34.95 Canaveral JSLI-2004-4703 20-Jun-04 North Cape 6.75 ± 0.00 6.73-6.80 34.91 ± 0.00 34.87-34.94 Canaveral
JSLI-2004-4704 21-Jun-04 South Cape 6.30 ± 0.00 6.28-6.36 34.90 ± 0.00 34.88-34.92 Canaveral JSLI-2004-4705 21-Jun-04 South Cape 6.29 ± 0.00 6.28-6.34 34.90 ± 0.00 34.89-34.92 Canaveral
17
Figure 6. Ship collected sonar tracks (top left) and resulting bathymetry maps (top right) from the deep coral area off Cape Lookout, NC (A). In this area additional data from our files were added for the bathymetry map. Bottom panel shows JSL submersible dive tracks in this area from 2000- 2005. All data are from Ross et al. (unpublished). See Fig. 1 to locate this area.
18
Figure 7. Ship collected sonar tracks (top left) and resulting bathymetry maps (top right) from the deep coral area off Cape Lookout, NC (B). Bottom panel shows JSL submersible dive tracks in this area from 2000-2005. All data are from Ross et al. (unpublished). See Fig. 1 to locate this area.
19
53 m over a distance of about 0.4 km. There is a small mound away from the main system (Fig. 7), and in general these mounds were less dramatic than those described above. They appeared to be of the same general construction as Bank A, built of coral rubble matrix that had trapped sediments. Extensive fields of coral rubble surrounded the area. Both living and dead corals were common on this bank, with some living bushes being quite large (Fig. 3).
Cape Fear Lophelia Bank
Aside from the map in EEZ-SCAN 87 Scientific Staff (1991) there are no published data from this coral mound and no indication that it was sampled before the studies initiated by Ross et al. (unpubl. data) between summer 2002 and fall 2005. Ross et al. located this bank based on estimated coordinates from the USGS survey (EEZ-SCAN 87 Scientific Staff 1991). As above, the JSL submersible was the major method for collecting bottom data on the reef proper. Nine dives were made on coral mounds in this area (Fig. 8, Table 1), and observations from these totaled about 20 hours. Sampling in this area was focused on a relatively small area (Fig. 8), but data are lacking to accurately estimate the size and area covered by coral mounds or rubble zones. Ross et al. JSL dives in this area ranged from 371-449 m (Table 1). Mean bottom temperatures ranged from 8.0 to 11.7 C, and as above mean bottom salinities were always near 35 (Table 2). These mounds rise nearly 80 m over a distance of about 0.4 km, and exhibit some of the most rugged habitat and vertical excursion of any area sampled (Fig. 4). This mound system also appears to be of the same general construction as Banks A and B, being built of coral rubble matrix with trapped sediments. Fields of coral rubble are common around the area. Both living and dead corals were common on this bank (Fig. 4).
CORAL BANKS OF THE BLAKE PLATEAU (South Carolina to Florida)
South of Cape Fear sediment/coral mounds are smaller and scattered; however, L. pertusa and other hard and soft corals populate the abundant hard substrates of the Blake Plateau in great numbers. Overall, species diversity of anthozoans and other associated sessile invertebrates (e.g., sponges, hydrozoans) increases south of Cape Fear, NC. For convenience, some deep coral study areas in this region have been named, giving the impression of isolated areas of coral habitat. It appears, however, that Blake Plateau coral habitats are larger and more continuous than these names imply. Future detailed mapping of the area (some planned for fall 2006) combined with ground truthing will clarify coral habitat distributions and the extent to which areas may require discrete names. There is existing research data for this area, but historically most of it was geological (see history above). Most deep coral expeditions south of North Carolina concentrated around the area described by Stetson et al. (1962), referred to as “Stetson Banks” (Fig. 9), an area off GA (“Savannah Banks”, Fig. 10), the Charleston Bump (Sedberry 2001), a large area straddling the GA- FL border (“Jacksonville Lithoherms”, Fig. 11) and numerous coral sites along the FL East coast (Figs. 12 and 13). General properties of these study areas were described in several papers by Reed and colleagues (Reed 2002, Reed unpubl. rept. to SAFMC 2004, Reed and Ross 2005, Reed et al. 2005, 2006). See the history section above for other references to this area. Because it is unclear that these coral study areas are physically separate, I do not discuss them
20
Figure 8. Ship collected sonar tracks (top left) and resulting bathymetry maps (top right) from the deep coral area off Cape Fear, NC. Bottom panel shows JSL submersible dive tracks in this area from 2000-2005. All data are from Ross et al. (unpublished). See Fig. 1 to locate this area and Fig. 4 for a 3-D view.
21
Figure 9. Ship collected sonar tracks (top left) and resulting bathymetry maps (top right) from the Stetson deep coral area off of SC. Bottom panel shows JSL submersible dive tracks in this area from 2000-2005. All data are from Ross et al. (unpublished). See Fig. 1 to locate this area.
22
Figure 10. Ship collected sonar tracks (top left) and resulting bathymetry maps (top right) from the Savannah Banks deep coral area off of SC-GA. Bottom panel shows JSL submersible dive tracks in this area from 2000-2005. All data are from Ross et al. (unpublished). See Fig. 1 to locate this area.
23
Figure 11. Ship collected sonar tracks (top left) and resulting bathymetry maps (top right) from the Jacksonville Banks deep coral area off of GA-FL. Bottom panel shows JSL submersible dive tracks in this area from 2000-2005. All data are from Ross et al. (unpublished). See Fig. 1 to locate this area.
24
Figure 12. Ship collected sonar tracks (top left) and resulting bathymetry maps (top right) from the deep coral area just north of Cape Canaveral, FL. Bottom panel shows JSL submersible dive tracks in this area from 2000-2005. All data are from Ross et al. (unpublished). See Fig. 1 to locate this area.
25
Figure 13. Ship collected sonar tracks (top left) and resulting bathymetry maps (top right) from the deep coral area just south of Cape Canaveral, FL. Bottom panel shows JSL submersible dive tracks in this area from 2000-2005. All data are from Ross et al. (unpublished). See Fig. 1 to locate this area.
26 individually. Note some differences between sites in the mapping and fish sections below. A few general observations are relevant. The Stetson Bank is a very large region of extremely diverse, rugged topography and bottom types. There is a deep canyon on the eastern side of this system with abundant corals on its western rim. While the surface waters of Stetson Bank are often outside the main Gulf Stream path, bottom currents can be quite strong. This is one of the deeper and more interesting of the Blake Plateau coral areas and warrants further exploration. The Savannah Bank system appears to have a heavier sediment load, perhaps because it is closest to the continental shelf. Deep corals occur there in scattered patches and are often less well developed than at other sites. Many sites in the “Jacksonville area” were composed of rocky ledges to which corals were attached, especially on the northern end. Bottom types in this area are diverse as is the fauna. Topographic highs, most having corals, are very abundant from the “Jacksonville area” to just south of Cape Canaveral (see also Reed et al. 2005, 2006). Faunal diversity is quite high in this region.
MAPPING DEEP CORAL BANKS
Basic SEUS study area maps were created by displaying varying combinations of data collected by the surface ships and submersible (S.W. Ross et al. Unpubl. data). The sonar track maps were simply the 2-D files of individual surveys color-coded by year with the addition of a scale, legend and north arrow (top left panel, Figs. 6-13). The 2-D raster files of the sonar data were combined with contours, labels, a scale, legend and north arrow to create the bathymetric maps (top right panels Figs. 6-13). The dive site maps were the various dive tracks, color-coded by year, laid over the 2-D raster files without the contours or the contour labels with addition of a scale bar, legend and north arrow (bottom panels Figs. 6-13). These base maps will be improved as additional data are analyzed, eventually leading to color-coded habitat maps with bathymetry. Three dimensional views will also be generated. An example of a three dimensional habitat map for one of the North Carolina sites is presented (Fig. 4, bottom).
Mapping Data Quality Issues Data available for this mapping effort varied greatly by year of the project. For instance, Knudsen sonar data were only available for two years (2004, 2005), and many sites have dive data from only one or two years. Data problems ranged from uncertain position information to missing dive track data. The maps generated with these data have some limitations. Site maps resulting from fewer sonar surveys or fewer dives display less details and may be less accurate. Data confirmation was difficult, but when dive or sonar data were available from multiple years at the same locations, the datasets did corroborate one another. These maps have been and will be used for planning research missions and displaying general habitat characteristics. They are good interim tools until more detailed mapping using multibeam sonar is undertaken. Such a survey of the area’s deep coral banks is sorely needed. Despite the above issues, it is important to note that these geospatial depth and habitat data represent the first such data for these areas of the SEUS slope. Most available maps are on large scales and/or present data at low resolution.
DEEP CORAL BANK FISH COMMUNITY DATA
Despite increasing research attention toward deep coral systems, knowledge of fish communities is still relatively lacking. In the cool temperate to boreal northeastern Atlantic Mortensen et al. (1995), Husebo et al. (2002), and Costello et al. (2005) noted that Lophelia habitat
27 seemed to be important to fishes. However, in the northwestern Atlantic Auster (2005) suggested that deep corals were no more important to fishes than other reef type habitats. Deep coral ecosystem fish data from the SEUS and Gulf of Mexico are more limited, with studies reporting only a few taxa, many not identified to species, from only a few areas (Messing et al. 1990; Wenner and Barans 2001; Reed et al. 2005, 2006). The summary below represents the first extensive treatment of fish communities on deep coral slope habitats of this region (Ross and Quattrini, ms in prep. a, b). We identified at least 57 unique taxa from our video analyses over all locations (2003-2004 data). A number of these species have never been reported from this region and some of those were thought to be rare (e.g., Caruso et al. in press). While most of the species richness was within prime reef or transition habitats (36 and 35 species, respectively) (Table 3, Fig. 14), the soft substrate off reef habitats supported a different but well developed fauna. The ichthyofauna of all three general habitat types was dominated by relatively few species, with little overlap in species between prime reef and off reef habitats. In particular, prime reef was characterized by Laemonema melanurum, Hoplostethus occidentalis, Beryx decadactylus, and Conger oceanicus. These species were never or only rarely observed on off reef, soft substrates and only rarely in the transition habitats. The off reef areas were characterized by L. barbatulum, Fenestraja plutonia, Myxine glutinosa, and Merluccius albidus, with F. plutonia and M. albidus never occurring on prime reef. When Helicolenus dactylopterus was observed away from reef habitat, it was usually near whatever structure was available (anemones, depressions). Transition habitat exhibited a mixture of species that could be found on either prime reef or off reef. The large, commercially important wreckfish (Polyprion americanus) seemed to move over several habitats from the base of mounds on rubble areas with little profile to the tops of ledges. Our preliminary conclusion from these data is that there is an obligate deep reef fish community that is tied to structured habitat (whether coral or rock). Ecologically, this parallels community structure found in shallow tropical reef systems. Species richness was higher at northern deep coral banks (off North Carolina) than those sampled from South Carolina to Florida. Results from multidimensional scaling analysis confirmed that regional differences existed in the ichthyofauna of the SEUS. The three North Carolina sites clustered together, the sites in the middle of the region (Stetson, Savannah, Jacksonville) grouped together, and the two Cape Canaveral areas grouped together. Similarity analysis further supported that these groups were significantly different from one another. The drivers of these assemblages (SIMPER analysis) were: NC Group - B. decadactylus, H. occidentalis, C. oceanicus, L. barbatulum, H. dactylopterus; Middle Group - L. melanurum, Nezumia sclerorhynchus, Trachyscorpia cristulata; Canaveral Group - N. sclerorhynchus, F. plutonia, Synaphobranchus kaupii . Additional analyses are in progress and will include additional years of data, especially from the Ross et al. 2005 cruise (Ross and Quattrini ms in prep. b). Hypotheses to be considered to explain these differences include: zoogeography effects (latitude/temperature), depth effects, habitat structure or quality influences, other (physical oceanography, food resources, recruitment).
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Table 3. Benthic fish species identified from analysis of Johnson-Sea-Link video data (2000-2004) at deep coral sampling locations along the southeastern United States slope from Cape Lookout, NC to just south of Cape Canaveral, FL. These are unpublished data of Ross et al. and are being analyzed for geographic and habitat patterns. Taxa Myxinidae Myxine glutinosa Chimaeridae Chimaera sp. Squalidae Cirrhigaleus asper Squalus cubensis Odontaspididae Odontaspis ferox Scyliorhinidae Scyliorhinus spp. Scyliorhinus meadi Scyliorhinus retifer Carcharhinidae Carcharhinus spp. Rajidae Dactylobatus armatus Fenestraja plutonia Mobulidae Manta birostris Synaphobranchidae Dysommina rugosa Synaphobranchus spp. Synaphobranchus kaupii Congridae Conger oceanicus Nettastomatidae Nettenchelys exoria Sternoptychidae Maurolicus weitzmani Polyipnus clarus Sternoptyx sp. Stomiidae Chauliodus sloani Chlorophthalmidae Chlorophthalmus agassizi Paralepididae Undetermined Myctophidae Undetermined Diaphus dumerilii
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Bythitidae Bellottia apoda Bythites gerdae Diplacanthopoma brachysoma Macrouridae Undetermined Nezumia spp. Nezumia aequalis Nezumia sclerorhynchus Moridae Laemonema spp. Laemonema barbatulum Laemonema melanurum Physiculus spp. Physiculus karrerae Merlucciidae Merluccius spp. Merluccius albidus Lophiidae Lophiodes beroe Lophiodes monodi Lophius cf. americanus Chaunacidae Chaunax stigmaeus Ogcocephalidae Dibranchus atlanticus Trachichthyidae Hoplostethus occidentalis Berycidae Beryx decadactylus Zeidae Zenopsis conchifera Scorpaenidae Helicolenus dactylopterus Idiastion kyphos Neomerinthe hemingwayi Pontinus rathbuni Trachyscorpia cristulata Acropomatidae Synagrops spp. Polyprionidae Polyprion americanus Serranidae Anthiinae Anthias woodsi Hemanthias aureorubens
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Trichiuridae Undetermined
Figure 14. Selected deep reef fishes from coral banks off the southeastern United States. From left to right and top to bottom: Helicolenus dactylopterus, Laemonema melanurum, Beryx decadactylus, Conger oceanicus, Hoplostethus occidentalis, Polyprion americanus. All photos from Ross et al. (unpubl. data).
31
SUMMARY AND RECOMMENDATIONS
These systems support a well developed community that appears to be different from the surrounding non-reef habitats. In fact, preliminary analyses suggest that the fish community on these deep reefs is composed of many species that do not (or at least rarely) occur off of the reefs. Therefore, they may be considered primary reef fishes, in a way similar to those on shallow reefs. Many fish species thought to be rare and/or outside their reported ranges have been found on these reefs (Ross et al. unpubl. data). Most likely these species only appeared to be rare because they occurred in a difficult to sample environment. Thus, these deep coral habitats support a fish community that appears to be tightly coupled to the habitat and has essentially escaped detection until recently. However, invertebrate associations with the reef habitat seem to be more opportunistic than is the case for certain fish species. Additional data are required from diverse habitats to confirm this. Some commercially-exploited deep-water fishes, like wreckfish (Polyprion americanus) (Vaughan et al. 2001) and blackbelly rosefish (Helicolenus dactylopterus), utilize Lophelia habitat extensively. Other potentially exploitable species are also associated with deep corals (royal red shrimps, rock crabs, bericiform fish species, eels). Signs of past fishing effort were observed on some coral banks, but the extent to which fishermen sample these areas is unknown. The potential for new deep water fisheries on and around these banks is unknown. The banks examined off of North Carolina are different than much of the coral habitat to the south on the Blake Plateau. The NC features are almost exclusively dominated by L. pertusa, the diversity of other corals being low. The fish and invertebrate faunas also differ between North Carolina and Blake Plateau deep coral areas.
Recommendations
Detailed mapping of the slope is critical to better understand these habitats and evaluate their contributions to slope ecology. Such mapping is the foundation for most other research and management activities. Multibeam mapping should be conducted as soon as possible, especially in the depth range of 350-500 m. While this recommendation relates to the whole slope of the SEUS, priority should be given to known coral sites and areas of suspected coral mounds.
Deeper areas of the Blake Plateau are virtually unexplored. The hard substrate region of the Blake Spur and Blake Escarpment and the 800-1000 m depth just to the West should be explored for deep coral habitat.
The HAPCs proposed by the SAFMC need some minor boundary adjustments based on additional recent data.
If protected areas are established for SEUS deep coral banks, long term monitoring and research plans should accompany this strategy.
Any deep water fisheries that currently exist or that develop on or near the deep coral banks should be carefully monitored and regulated as deep water fauna are highly vulnerable to over fishing and the habitat is subject to permanent destruction.
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Of the many important ecological/biological studies that could be proposed, a broad trophodynamics study of the coral banks and surrounding area (whole water column) would probably provide the most impact for funds expended. Knowing the flow of energy in a system facilitates evaluation of anthropogenic impacts and the allows predictions about the consequences of natural change.
Life history data for most deep water species are lacking, and this data need should be addressed.
Biodiversity of deep coral habitats requires additional study as these habitats appear to host huge numbers of species.
A regional working group composed of scientists and relevant agency personnel should be formed to begin evaluating data, deep reef status, and to take the lead on formulation of plans to study and manage these habitats.
ACKNOWLEDGMENTS
NOAA Office of Ocean Exploration (grants NA16RP2696, NA030AR4600090, NA040AR4600056 to S.W. Ross, lead PI) largely supported field work and some of the data analyses for this research. United States Geological Survey (through the State Partnership Program), Minerals Management Service, and South Atlantic Fishery Management Council contributed funds to help with analyses. USGS Florida Integrated Science Center provided personnel and logistics support for most field operations. Dr. M.S. Nizinski provided assistance with all aspects of invertebrate data. A. Quattrini led many aspects of data organization and analyses of fish community data. I thank M. Partyka for all GIS work and production of various maps.
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DEEP-WATER CORAL REEFS OF FLORIDA, GEORGIA AND SOUTH CAROLINA: A SUMMARY OF THE DISTRIBUTION, HABITAT, AND ASSOCIATED FAUNA
by John K. Reed Harbor Branch Oceanographic Institution 5600 U.S. 1, North, Fort Pierce, FL 34946 Phone- 772-465-2400 x205, Fax- 772-461-2221 Email- [email protected]
Contract No: SA-04-05-NC/UNCW Submitted to: South Atlantic Fishery Management Council One Southpark Circle, Suite 306 Charleston, SC 29407
[All rights reserved. Authorization requested by the author for photocopying or electronic distribution of any parts of this document. Copying or electronic distribution of tables or figures must include accompanying caption with complete citation.]
October 20, 2004
ABSTRACT
This report was compiled at the request of the South Atlantic Fishery Management Council (SAFMC) to provide a preliminary, general summary on the status of current knowledge concerning deep-water (> 200 m) reefs off the southeastern U.S. from Florida to North Carolina. The outcome will provide target areas of deep-water, live-bottom habitats for: 1) potential designation as Habitat Areas of Particular Concern (HAPC) or Marine Protected Areas (MPA) by the SAFMC, and 2) high-resolution habitat maps and habitat characterization studies. The resource potential of the deep-water habitats in this region is unknown in terms of fisheries and novel compounds yet to be discovered from associated fauna that may be developed as pharmaceutical drugs. Although these habitats have not been designated as MPAs or HAPCs, they are incredibly diverse and irreplaceable resources. Activities involving bottom trawling, pipelines, or oil/gas production could negatively impact these reefs. This report primarily summarizes recent submersible data regarding deep-water reefs off Florida but also includes sites off Georgia and South Carolina. A report on the North Carolina reefs has been submitted separately by Dr. Steve Ross, UNCW. This report does not include the deep-water Oculina reefs off central eastern Florida or deep shelf-edge reefs with hermatypic coral (<100 m). The sites included in this report are the following: 1) Stetson Reefs- hundreds of pinnacles along the eastern Blake Plateau off South Carolina include a 152-m tall pinnacle (822 m depth) where recent submerisible dives discovered live bushes of Lophelia coral, sponges, gorgonians, and black coral bushes. 2) Savannah Lithoherms- numerous lithoherms at depths of 550 m with relief up to 60 m provide live-bottom habitat. 3) East Florida Lophelia Reefs- echosounder transects along a 222-km stretch off eastern Florida (depth 700-800 m) mapped hundreds of 15- 152 m tall coral pinnacles and lithoherms. 4,5) Miami Terrace and Pourtales Terrace- Miocene age terraces off southeastern Florida and the Florida reef tract provide high-relief, hard-bottom habitats and rich benthic communites. 6) SW Florida Lithoherms- in the Gulf of Mexico off the southwestern Florida shelf slope, 15-m tall Lophelia coral lithoherms (500 m depth) are described the first time from SEABEAM and ROV dives.
JUSTIFICATION
The South Atlantic Fishery Management Council (R. Pugliese) requested that this preliminary summary report on the state of knowledge of Deep Sea Coral Ecosystems (DSCE) in the region be available in time for the Habitat Advisory Panel meeting of the SAFMC, October 26, 2004. The Council needs immediate scientific data and maps as it considers designation of new Habitat Areas of Particular Concern (HAPC) to protect DSCE areas. Such protection may be needed to prevent long-term (perhaps permanent) damage, such as has occurred on shallower Oculina reefs off Florida and Lophelia banks in the northeastern Atlantic, both destroyed in part by trawling. After trawlers were banned from the Oculina HAPC, there is justified concern that trawlers may move to deeper habitats in search of valuable commercial fisheries, such as royal red shrimp or benthic finfish. NOAA is currently developing priority mapping sites, including Marine Protected Areas and DSCE. NOAA OE funding for 2005 will likely support habitat mapping of shelf-edge and deep-water reef habitats in the South Atlantic Bight and Gulf of Mexico. Data compiled in this report provides potential targets for future mapping, MPAs and HAPCs. The
2 resource potential of the deep-water habitats in this region is unknown in terms of fisheries and novel compounds yet to be discovered from associated fauna that may be developed as pharmaceutical drugs. Although these habitats are not currently designated as MPAs or HAPCs, they are incredibly diverse and irreplaceable resources. Activities involving bottom trawling, pipelines, or oil/gas production could negatively impact these reefs.
OBJECTIVES
Objectives of this report and accompanying DVD are the following: 1) Compile list of references regarding geology and biology of deep-water reef habitats in the South Atlantic Bight, Straits of Florida and southwest Florida slope. 2) Describe general habitat for each reef type and region (northeastern Florida, Straits of Florida, southwest Florida slope, and areas of DSCE off Georgia and South Carolina). 3) Provide representative digital still images and video clips for examples of reef types and regions (on DVD). 4) Provide species list of dominant benthic invertebrates that are directly associated with these reefs based on recent collections and observations by the PI (based on current status of taxonomic identifications). 5) Provide species list of fish that are directly associated with these reefs based on recent collections and observations by the PI (based on current status of taxonomic identifications). 6) Provide general maps of known DSCE reefs in the region.
BACKGROUND
Deep-water reefs are sometimes defined as bioherms, coral banks, or lithoherms (Teichert, 1958; Stetson et al., 1962; Neumann et al., 1977; Wilson, 1979; Reed, 1980; Freiwald et al. 1997; Fosså et al. 2000; Paull et al., 2000). Some deep-water reefs consist of caps of living coral on mounds of unconsolidated mud and coral debris, such as some Oculina and Lophelia coral reefs (Reed 2002a,b), whereas deep-water lithoherms are defined as high-relief, lithified carbonate limestone mounds rather than unconsolidated mud mounds (Neumann et al., 1977). Rogers (1999) has suggested that deep-water coral bioherms fall within the definition of a coral reef based on their physical and biological characteristics. Various types of deep-water, high-relief bioherms are common off the southeastern United States, along the base of the Florida-Hatteras Slope, on the Blake Plateau, in the Straits of Florida, and eastern Gulf of Mexico. Only a small percentage of deep-water reefs have had their benthic and fish resources characterized.
Recent research expeditions by Principal Investigator (PI), J. Reed, Harbor Branch Oceanographic Institution (HBOI), using HOVs (human occupied vehicle) and ROVs (remotely operated vehicle) along with previous research by the PI in the 1990s and 1980s, have compiled new information on the status, distribution, habitat, and biodiversity of some of these relatively unknown and newly discovered deep reef ecosystems. In 2004, during a State of Florida funded mission with the Johnson-Sea-Link (JSL) Submersible, the PI discovered nearly 300 potential targets during echosounder transects that may be newly discovered deep-water reefs off the east coast of Florida, some of which are up to 168 m (550 feet) in height at depths of 732 m (2400 feet) (Reed and Wright, 2004; Reed et al., 2004b). Expeditions in 2002 and 2003 for biomedical
3 research by the PI and funded by the National Oceanic and Atmospheric Administration’s Office of Ocean Exploration (NOAA OE) enabled preliminary exploration of additional deep-water reef sites in the western Atlantic (Blake Plateau) and eastern Gulf of Mexico on southwest Florida shelf slope (Reed, 2003, 2004; Reed and Pomponi, 2002b; Reed et al., 2002, 2003, 2004d). These were the first HOV and ROV dives ever to document the habitat and benthic biodiversity of some of these relatively unknown deep-water reefs. A small scale, high-definition topographic SEABEAM map was also conducted by the PI at the southwest Florida site. Considerable work remains to analyze these data and prepare for scientific publications (three papers in preparation or submitted by PI: Florida’s Deep-Water Lophelia Reefs; Miami Terrace Deep-Water Reefs; Deep-Water Sinkholes and Bioherms of Pourtales Terrace- Habitat and Biology). These are very preliminary analyses based on only a few submersible or ROV dives at the various sites.
Florida DSCE Deep sea coral ecosystems (DSCE) in U.S. EEZ waters exist along the eastern and southwest Florida shelf slope (in addition to the Oculina Marine Protected Area and deep shelf-edge reefs with hermatypic coral). These include a variety of high-relief, hard-bottom, live-bottom habitats at numerous sites along the base of the Florida-Hatteras Slope off northeastern and central eastern Florida, the Straits of Florida, the Miami Terrace and Pourtales Terrace off southeastern Florida, and the southwestern Florida shelf slope. The predominate coral on these reefs are the azooxanthellate, colonial scleractinian corals, Lophelia pertusa, Madrepora oculata, and Enallopsammia profunda; various species of hydrocorals of the family Stylasteridae, and species of the bamboo octocoral of the family Isididae. Various types of high-relief, live-bottom habitat have been discovered in the area: Lophelia mud mounds, lithoherms, sinkholes, ancient Miocene escarpments and karst topographic features (Reed 2002b; Reed et al., 2004a,b). These all provide hard-bottom substrate and habitat for sessile macrofauna including deep-water corals, octocorals (gorgonians), black coral, and sponges, which in turn provide habitat and living space for a relatively unknown but biologically rich and diverse community of associated fish, crustaceans, mollusks, echinoderms, polychaete and sipunculan worms, and other macrofauna, many of which are undoubtedly undescribed species. Our preliminary studies have found new species of octocorals and sponges from some these sites (Reed et al., 2004 a,b).
RESULTS
Coral Description and Distribution (from Reed, 2002a) The dominant colonial scleractinian coral species forming deep-water reefs in the western North Atlantic region are Oculina varicosa, Lophelia pertusa, and Enallopsammia profunda, although other branching colonial scleractinia may also occur, including Solenosmilia variabilis and Madrepora oculata (Figs. 1 and 2). Numerous solitary coral species are also common (Cairns, 1979).
Lophelia pertusa (Linnaeus, 1758) (= L. prolifera): This coral forms massive, dendroid, bushy colonies, 10-150 cm in diameter, with anastomosing branches (Figure 1). Its distribution ranges in the western Atlantic from Nova Scotia to Brazil and the Gulf of Mexico, and also in the eastern Atlantic, Mediterranean, Indian, and eastern Pacific Oceans at depths of 60-2170 m (Cairns, 1979). Along with Enallopsammia profunda, it is the primary constituent of deep-water reefs at the base of
4 Figure 1. Coral colony and branch tip: top- Oculina varicosa (80m); middle- Lophelia pertusa (490m); bottom- Enallopsammia profunda (585m). (scale lines = 1 cm; top left fig. Scale = 5 cm) (from Reed, 2002a; Hydrobiologia 471: 57-69)
5 the Florida-Hatteras slope and at depths of 500-800 m from Miami to South Carolina (Figure 3, Region B and C). In addition, over 200 banks have been mapped at depths of 640-869 m (Region D) on the outer eastern edge of the Blake Plateau (Stetson et al., 1962; Popenoe and Manheim, 2001). Elsewhere deep-water Lophelia reefs are known from the Gulf of Mexico (Ludwick & Walton, 1957; Moore & Bullis, 1960; Newton et al., 1987) and the eastern Atlantic off Norway and Scotland (Teichert, 1958; Wilson, 1979a; Mortensen et al., 1995; Freiwald et al., 1997, 1999). In the eastern Atlantic, Madrepora oculata commonly occurs with Lophelia rather than E. profunda.
Enallopsammia profunda (Pourtalès, 1867) (=Dendrophyllia profunda): This species also forms dendroid, massive colonies up to 1 m in diameter (Figure 1). It is endemic to the western Atlantic and ranges from the Antilles in the Caribbean to Massachusetts at depths of 146-1748 m (Cairns, 1979). E. profunda occurs with L. pertusa at Regions B, C, and D (Figure 3). It appears to be the primary constituent of the deep-water reefs at Site D except at the tops of the mounds where L. pertusa is more prevalent (Stetson et al., 1962).
Six regions (B-D, G-I) of deep-water reef habitats off southeastern U.S. from Florida to South Carolina may be considered targets for potential HAPCs (Figs. 3-8). Figure 3 shows the general boundaries of Regions A-H off eastern Florida, Georgia, and South Carolina. It also includes the Oculina Reefs (A) that are already designated as an HAPC and two regions (E,F) that are within Bahamian waters, but are not discussed in this report (see Reed 2002a,b). Recent submersible dive sites and echosounder locations of high-relief reef habitat off the east coast are shown in Figure 4 (see Table 1 for corresponding dive sites). Details of the Lophelia mounds of Region D (Stetson’s Reefs) are shown in Figure 5 (Popenoe and Mannheim, 2001). Figure 6 shows the bathymetry and submersible dive sites at Region G, Miami Terrace Escarpment. Figure 7 shows the bathymetry and submersible dive sites at Region H, Pourtales Terrace. Figure 8 shows the bathymetry and ROV dive sites in the Gulf of Mexico at Region I, Southwest Florida Lithoherms.
Figure 2. Depth range and maximum relief of deep-water coral reefs off southeastern U.S.A. Dominant colonial coral listed for each site (see Figure 3 for site locations). (from Reed, 2002a; Hydrobiologia 471: 57-69)
6 Figure 3. Deep-water coral reef regions off southeastern U.S.A. (see Table 1 for locations). ?= Johnson-Sea-Link I and II submersible dive sites and echosounder sites of high-relief reefs; Regions: A=Oculina Coral Reefs, B= East Florida Lophelia Reefs, C= Savannah Lophelia Lithoherms, D= Stetson’s Reefs (D1= region of dense pinnacles), E= Enallopsammia Reefs (Mullins et al., 1981), F= Bahama Lithoherms (Neumann et al., 1977), G= Miami Terrace Escarpment. (from Reed et al., 2004b; chart from NOAA, NOS, 1986)
7 Figure 4. Submersible dive sites and echosounder sites on deep-water reefs off southeastern U.S.A. (see Table 1 for locations). ?# = Johnson-Sea-Link I and II submersible dive sites, F# = high-relief pinnacles from echosounder transects. (from Reed et al., 2004b; chart from NOAA, NOS, 1986)
8 Figure 5. Detailed chart of high-relief region with Lophelia coral mounds on Charleston Bump, Blake Plateau. (from Popenoe and Manheim, 2001; American Fisheries Society Symposium 25: 43- 94)
9 Figure 6. Bathymetry and submersible dive sites on Miami Terrace Escarpment at Region G (see Table 1 for locations). ?= Johnson-Sea-Link I submersible dive sites. (from Reed et al., 2004b; chart from Ballard and Uchupi, 1971; MTS Journal 5: 43-48)
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Figure 7. Bathymetry and submersible dive sites on Pourtalès Terrace at Region H (see Table 2 for locations). ?= Johnson-Sea-Link and Clelia submersible dive sites; JS= Jordan Sinkhole, MS= Marathon Sinkhole, TB1= Tennessee Humps Bioherm #1, TB2= Tennessee Humps Bioherm #2, AB3= Alligator Humps Bioherm #3, AB4= Alligator Humps Bioherm #4. (from Reed et al., 2004b; chart from Malloy and Hurley, 1970; Geol. Soc. Amer. Bull. 81: 1947-1972)
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Figure 8. Deep-water coral lithoherms and ROV dive sites at Region I off southwest Florida slope (see Table 1 for locations). ?= Innovator ROV dive sites. (from Reed et al., 2004b; chart from NOAA, NOS, 1986)
12 Deep-water Coral Reef Communities (from Reed, 2002a,b) The deep-water coral reefs support very rich communities of associated invertebrates. Faunal diversity on the Oculina reefs is equivalent to many shallow-water tropical reefs. Over 20,000 individual invertebrates were found living among the live and dead branches of 42 small Oculina colonies from deep and shallow water, yielding 230 species of mollusks, 50 species of decapods, 47 species of amphipods, 21 species of echinoderms, 15 species of pycnogonids, and numerous other taxa (Reed et al., 1982; Reed & Hoskin, 1987; Reed & Mikkelsen, 1987; Child, 1998). A striking difference between the Oculina and Lophelia reefs is that larger sessile invertebrates such as massive sponges and gorgonians are common on the Lophelia reefs but are not common on the deep-water Oculina reefs. The coral itself is a dominant component providing habitat on both the Oculina and Lophelia reefs. The percentage of live coral coverage is generally low on the majority of Lophelia and Oculina reefs in this region (1-10%); however, some areas may have nearly 100% live cover and some areas may have extensive areas of 100% dead coral rubble.
In comparison, Rogers’ (1999) review of literature on deep-water Lophelia coral reefs in the northeastern Atlantic recorded 886 species of associated invertebrates. Quantified analyses of live and dead colonies of Lophelia pertusa from the Faeroe shelf off of Scotland resulted in 298 species, dominated by Polychaeta (67 sp.), Bryozoa (45 sp.), Mollusca (31 sp.), Porifera (29 types), and Crustacea (15 sp.) (Jensen & Frederiksen, 1992). Studies of infauna associated with the Lophelia reefs of the western Atlantic reefs off North Carolina have just begun (Ross, in prep).
Region D: Stetson Reefs, Eastern Blake Plateau (from Reed, 2002a; Reed et al., 2004b) This site is on the outer eastern edge of the Blake Plateau, ~120 nm SE of Charleston, South Carolina, at depths of 640-869 m (Table 1, Figs. 3-5). Over 200 coral mounds up to 146 m in height occur over this 6174 km2 area that was first described by Thomas Stetson from echo soundings and bottom dredges (Stetson et al., 1962; Uchupi, 1968). These were described as steep-sloped structures with active growth on top of the banks. Live coral colonies up to 50 cm in diameter were observed with a camera sled. E. profunda (=D. profunda) was the dominant species in all areas although L. pertusa was concentrated on top of the mounds. Densest coral growth occurred along an escarpment at Region D1. Stetson et al. (1962) reported an abundance of hydroids, alcyonaceans, echinoderms, actiniaria, and ophiuroids, but a rarity of large mollusks. The flabelliform gorgonians were also current-oriented. Popenoe and Manheim (2001) have made detailed geological maps of this Charleston Bump region which also indicate numerous coral mounds (Fig. 5).
Recent fathometer transects by the PI indicated dozens and possibly hundreds of individual pinnacles and mounds within the small region that we surveyed which is only a fraction of the Stetson Bank area (Reed and Pomponi, 2002b; Reed et al., 2002; Reed et al., 2004b). From our fathometer transects, two pinnacle regions were selected. Three submersible dives were made on “Pinnacle 3” and four dives on “Stetson’s Peak” which is described below (Table 1). A small subset of the Stetson Bank area was first mapped by six fathometer transects covering ~28 nm2 (6 nm x 4.7 nm; 31o59.03’N to 32o05.03’N and 77o42.75’W to 77o37.98’W), in which six major peaks or pinnacles and four major scarps were plotted. The base depth of these pinnacles ranged from 689 m to 643 m, with relief of 46 to 102 m. A subset of this was further mapped with 70 fathometer transects spaced 250 m apart (recording depth, latitude and longitude ~ every 3
13 seconds), covering an area of 1 x 1.5 nm (32o00.5’N to 32o01.5’N and 77o40.0’W to 77o42.5’W), resulting in a 3-D bathymetric GIS Arcview map of a major feature, which we named Stetson’s Pinnacle (Fig. 9).
Figure 9. Echosounder profile of Stetson’s Pinnacle (depth 780 m, relief 153 m). (from Reed et al., 2004b) -1500 feet
500 ft tall Lithoherm 1 nm N-S
-2700 feet
Stetson’s Pinnacle was 780 m at the south base and the peak was 627 m (differential GPS coordinates of submersible at the peak: 32o01.6882’N, 77o39.6648’W). This represents one of the tallest Lophelia coral lithoherms known, nearly 153 m in relief. The linear distance from the south base to the peak was ~0.5 nm. The lower flank of the pinnacle from ~762 m to 701 m on the south face was a gentle slope of 10-30o with a series of 3-4 m high ridges and terraces that were generally aligned 60-240o across the slope face. These ridges were covered with nearly 100% Lophelia coral rubble, 15-30 cm colonies of live Lophelia, and standing dead colonies of Lophelia, 30-60 cm tall. Very little rock was exposed, except on the steeper exposed, eroded faces of the ridges. Some rock slabs, ~30 cm thick, have slumped from these faces. From 701 m to 677 m the slope increased from ~45o to 60o. From 671 m to the peak, the geomorphology was very complex and rugged, consisting of 60-90o rock walls and 3-9 m tall rock outcrops. Colonies of Lophelia, 30-60 cm tall, were more common, and some rock ledges had nearly 100% cover of live Lophelia thickets. The top edge of the pinnacle was a 30 cm thick rock crust which was undercut from erosion; below this was a 90o escarpment of 3-6 m. The peak was a flat rock plateau at 625- 628 m and was approximately 0.1 nm across on a S-N submersible transect. The north face was not explored in detail but is a vertical rock wall from the peak to ~654 m then grades to a 45o slope with boulders and rock outcrops.
Dominant sessile macrofauna consisted of scleractinia, stylasterine hydrocorals, gorgonacea and sponges (Table 3). The colonial scleractinia were dominated by colonies of Lophelia pertusa (30-60 cm tall) and Enallopsammia profunda, and Solenosmilia variabilis were present. Small
14 stylasterine corals (15 cm tall) were common and numerous species of solitary cup corals were abundant. Dominant octocorallia consisted of colonies of Primnoidae (15-30 cm tall), paramuriceids (60-90 cm), Isididae bamboo coral (15-60 cm), stolonifera, and stalked Nephtheidae (5-10 cm). Dominant sponges consisted of Pachastrellidae (25 cm fingers and 25- 50 cm plates), Corallistidae (10 cm cups), Hexactinellida glass sponges (30 cm vase), Geodia sp. (15-50 cm spherical), and Leiodermatium sp. (50 cm frilly plates). Although motile fauna were not targeted, some dominant groups were noted. No large decapods crustaceans were common although some red portunids were observed. Two species of echinoids were common, one white urchin and one stylocidaroid. No holothurians or asteroids were noted. Dense populations of Ophiuroidea were visible in close-up video of coral clusters and sponges. No large Mollusca were noted except for some squid. Fish consisted mostly of benthic gadids and rattails. On the steeper upper flank, from 671 to 625 m the density, diversity, and size of sponges increased; 15- 50 cm macro sponges were more abundant. Massive Spongosorites sp. were common, Pachastrellidae tube sponges were abundant, and Hexactinellida glass sponges were also common. On the peak plateau the dominant macrofauna were colonies of Lophelia pertusa (30- 60 cm tall), coral rubble, Phakellia sp. fan sponges (30-50 cm), and numerous other demosponges were abundant. No large fish were seen on top.
Region C: Savannah Lithoherms, Blake Plateau (from Reed, 2002a; Reed et al., 2004b) A number of high-relief lithoherms occur within this region of the Blake Plateau, ~90nm east of Savannah, Georgia (Table 1; Figs. 3,4). Region C is at the base of the Florida-Hatteras Slope, near the western edge of the Blake Plateau, and occurs in a region of phosphoritic sand, gravel and rock pavement on the Charleston Bump (Sedberry, 2001). Wenner and Barans (2001) described 15-23 m tall coral mounds in this region that were thinly veneered with fine sediment, dead coral fragments and thickets of Lophelia and Enallopsammia. They found that blackbellied rosefish and wreckfish were frequent associates of this habitat. In general, the high-relief Lophelia mounds occur in this region at depths of 490-550 m and have maximum relief of 61 m (Table 1). JSL-II dives 1690, 1697 and 1698 reported a coral rubble slope with <5% cover of 30 cm, live coral colonies (Reed, 2002a). On the reef crest were 30-50 cm diameter coral colonies covering ~10% of the bottom. Some areas consisted of a rock pavement with a thin veneer of sand, coral rubble, and 5-25 cm phosphoritic rocks. At Alvin dive sites 200 and 203, Milliman et al. (1967) reported elongate coral mounds, approximately 10 m wide and 1 km long, that were oriented NNE-SSW. The mounds had
25-37o slopes and 54 m relief. Live colonies (10-20 cm diameter) of E. profunda (=D. profunda) dominated and L. pertusa (=L. prolifera) was common. No rock outcrops were observed. These submersible dives found that these lithoherms provided habitat for large populations of massive sponges and gorgonians in addition to the smaller macroinvertebrates which have not been studied in detail. Dominant macrofauna included large plate-shaped sponges (Pachastrella monilifera) and stalked, fan-shaped sponges (Phakellia ventilabrum), up to 90 cm in diameter and height. At certain sites (JSL-II dive 1697), these species were estimated at 1 colony/10 m2. Densities of small stalked spherical sponges (Stylocordyla sp., Hadromerida) were estimated in some areas at 167 colonies/10 m2. Hexactinellid (glass) sponges such as Farrea? sp. were also common. Dominant gorgonacea included Eunicella sp. (Plexauridae) and Plumarella pourtalessi (Primnoidae).
Recent fathometer transects by the PI at Savannah Lithoherm Site #1 (JSL II-3327) extended 2.36 nm S-N (31o40.3898’N to 31o42.7558’N along the longitude of 79o08.5’W) revealed a massive lithoherm feature that consisted of five major pinnacles with a base depth of 549 m,
15 minimum depth of 465 m, and maximum relief of 83 m (Reed and Pomponi, 2002b; Reed et al., 2002; Reed et al., 2004b). The individual pinnacles ranged from 9 to 61 m in height. A single submersible transect, south to north, on Pinnacle #4 showed a minimum depth of 499 m. The south flank of the pinnacle was a gentle 10-20o slope, with ~90% cover of coarse sand, coral rubble and some 15 cm rock ledges. The peak was a sharp ridge oriented, NW-SE, perpendicular to the prevailing 1 kn current. The north side face of the ridge was a 45o rock escarpment of about 3 m which dropped onto a flatter terrace. From a depth of 499 to 527 m, the north slope formed a series of terraces or shallow depressions, ~9-15 m wide, that were separated by 3 m high escarpments of 30-45\o. Exposed rock surfaces showed a black phosphoritic rock pavement. The dominant sessile macrofauna occurred on the exposed pavement of the terraces and in particular at the edges of the rock outcrops and the crest of the pinnacle. The estimated cover of sponges and gorgonians was 10% on the exposed rock areas. Colonies of Lophelia pertusa (15-30 cm diameter) were common but not abundant with ~1% coverage. Dominant Cnidaria included several species of gorgonacea (15-20 cm tall), Primnoidae, Plexauridae (several spp.), Antipathes sp. (1 m tall), and Lophelia pertusa (Table 3). Dominant sponges included large Phakellia ventilabrum (fan sponges, 30-90 cm diameter), Pachastrellidae plate sponges (30 cm), Choristida plate sponges (30 cm), and Hexactinellid glass sponges. Motile fauna consisted of decapod crustaceans (Chaceon fenneri, 25 cm; and Galatheidae, 15 cm) and mollusks. Few large fish were observed but a 1.5 m swordfish, several 1 m sharks, and numerous blackbelly rosefish were noted.
A fathometer transect by the PI at Savannah Lithoherm Site 2 extended 4.6 nm, SW to NE (31o42.0812’N, 79o07,6333’W to 31o45.5025’N, 79o04.0797’W), mapped 8 pinnacles with maximum depth of 549 m and relief of 15-50 m (Fig. 10).
Figure 10. Echosounder profile of Savannah Lithoherm, Site 2, Pinnacle #1 (depth 537 m, relief 50 m). (from Reed et al., 2004b)
Savannah Lithoherm
150 ft tall lithoherm
-1600 feet
-1800 feet
Submersible dives were made on Pinnacles 1, 5 and 6 of this group (Table 1). Pinnacle 1 was the largest feature of this group; the base was 537 m and the top was 487 m. The south face, from a depth of 518 to 510 m, was a gentle 10o slope, covered with coarse brown sand and
16 Lophelia coral rubble. A 3-m high ridge of phosphoritic rock, extended NE-SW, cropped out at a depth of 510 m. This was covered with nearly 100% cover of 15 cm thick standing dead Lophelia coral and dense live colonies of Lophelia pertusa (15-40 cm). From depths of 500 m to 495 m were a series of exposed rock ridges and terraces, that were 3-9 m tall with 45o slopes. Some of the terraces were ~30 m wide. Each ridge and terrace had thick layers of standing dead Lophelia, and dense live coral. These had nearly 100% cover of sponges (Phakellia sp., Geodia sp., Pachastrellidae, and Hexactinellida), scleractinia (Lophelia pertusa, Madrepora oculata), stylasterine hydrocorals, numerous species of gorgonacea (Ifalukellidae, Isididae, Primnoidae), and 1 m bushes of black coral (Antipathes sp.). Deep deposits of sand and coral rubble occurred in the depressions between the ridges. The north face, from 500 m to 524 m was a gentle slope of 10o, that had deep deposits of coarse brown foraminiferal sand and coral rubble. Exposed rock pavement was sparse on the north slope, but a few low rises with live bottom habitat occurred at 524 m. Dominant mobile fauna included decapod crustaceans (Chaceon fenneri, 15 cm Galatheidae), rattail fish, and 60 cm sharks were common.
Region B: Florida Lophelia Pinnacles (from Reed, 2002a; Reed et al., 2004b) Numerous high-relief Lophelia reefs and lithoherms occur in this region at the base of the Florida- Hatteras Slope and at depths of 670-866 m (Table 1, Figs. 3, 4). The reefs in the southern portion of this region form along the western edge of the Straits of Florida and are 15-25 nmi east of the Oculina coral banks Marine Protected Area (MPA). Along a 222-km stretch off northeastern and central Florida (from Jacksonville to Jupiter), nearly 300 mounds from 8 to 168 m in height (25- 550 ft) were recently mapped by the PI using a single beam echosounder (Fig. 11; Reed et al., 2004b). Between 1982 and 2004, dives with the Johnson-Sea-Link (JSL) submersibles and ROVs by the PI confirmed the presence of Lophelia mounds and lithoherms in this region (Reed, 2002a; Reed et al., 2002; Reed and Wright, 2004; Reed et al., 2004b). The northern sites off Jacksonville and southern Georgia appeared to be primarily lithoherms which are pinnacles capped with exposed rock (described in part by Paull et al., 2000), whereas the features from south of St. Augustine to Jupiter were predominately Lophelia coral pinnacles or mud mounds capped with dense 1-m-tall thickets of Lophelia pertusa and Enallopsammia profunda with varying amounts of coral debris and live coral. Dominant habitat-forming coral species were Lophelia pertusa, Madrepora oculata, Enallopsammia profunda, bamboo coral (Isididae), black coral (Antipatharia), and diverse populations of octocorals and sponges (Reed et al., 2004b).
Paull et al. (2000) estimated that over 40,000 coral lithoherms may be present in this region of the Straits of Florida and the Blake Plateau. Their dives with the Johnson-Sea-Link submersible and the U.S. Navy’s submarine NR-1 described a region off northern Florida and southern Georgia of dense lithoherms forming pinnacles 5 to 150 m in height with 30-60o slopes that had thickets of live ahermatypic coral (unidentified species, but photos suggest Lophelia and/or Enallopsammia). The depths range from 440 to >900 m but most mounds were within 500-750 m. Each lithoherm was ~100-1000 m long and the ridge crest was generally oriented perpendicular to the northerly flowing Gulf Stream current (25-50 cm s-1 on flat bottom, 50-100 cm s-1 on southern slopes and crests). Thickets of live coral up to 1 m were mostly found on the southern facing slopes and crests whereas the northern slopes were mostly dead coral rubble. These were termed lithoherms since the mounds were partially consolidated by a carbonate crust, 20-30 cm thick, consisting of micritic wackestone with embedded planktonic foraminifera, pteropods, and coral debris (Paull et al., 2000).
17 Figure 11. Height of Lophelia pinnacles and lithoherms on echosounder transects from Jacksonville to Jupiter, Florida at depths of 600 to 800 m. (from Reed et al., 2004b)
All Pinnacles
200 150 100 50 Height (m) 0 1 17 33 49 65 81 97 113 129 145 161 177 193 209 225 241 257 273 Individual Lophelia Pinnacles
A recent echosounder transect by the PI revealed a massive lithoherm, 3.08 nm long (N-S) that consisted of at least 7 individual peaks with heights of 30-60 m (Fig. 12; Reed and Wright, 2004; Reed et al., 2004b). The maximum depth was 701 m with total relief of 157 m. Three submersible dives (JSL II-3333, 3334; I-4658) were made on Peak 6 of pinnacle #204B (30o30.1194’N, 79o39.4743’W) which was the tallest individual feature of the lithoherm with maximum relief of 107 m and a minimum depth at the peak of 544 m (Reed et al., 2004b). The east face was a 20-30o slope and steeper (50o) near the top. The west face was a 25-30o slope which steepened to 80o from 561 m to the top ridge. The slopes consisted of sand and mud, rock pavement and rubble. A transect up the south slope reported a 30-40o slope with a series of terraces and dense thickets of 30-60 cm tall dead and live Lophelia coral that were mostly found on top of mounds, ridges and terrace edges. One peak at 565 m had dense thickets of live and dead standing Lophelia coral (~20% live) and outcrops of thick coral rubble. Dominant sessile fauna consisted of Lophelia pertusa, abundant Isididae bamboo coral (30-60 cm) on the lower flanks of the mound, Antipatharia black coral, and abundant small octocorals including the gorgonacea (Placogorgia sp., Chrysogorgia sp, and Plexauridae) and Nephtheidae soft corals (Anthomastus sp., Nephthya sp.). Dominant sponges consisted of Geodia sp., Phakellia sp., Spongosorites sp. Petrosiidae, Pachastrellidae, and Hexactinellida (Table 3).
Further south off Cape Canaveral, echosounder transects by the PI on Lophelia Pinnacle #113 (28o47.6258’N, 79o37.5859’W) revealed a 61 m tall pinnacle with maximum depth of 777 m (Table 1; Fig. 13). The width (NW-SE) was 0.9 nm and consisted of at least 3 individual peaks or ridges on top, each with 15-19 m relief. One submersible dive (JSL II-3335) reported 30-60o slopes, with sand, coral rubble, and up to 10% cover of live coral. No exposed rock was observed. This appeared to be a classic Lophelia mud mound.
18 Figure 12. Echosounder profile of Jacksonville Lithoherm, Pinnacle #204B (depth 701 m, relief 157 m). (from Reed et al., 2004b)
-1800 feet
400 ft tall Lithoherm (3 nm N-S 1 nm E-W)
-2500 feet
Figure 13. Echosounder profile of Cape Canaveral Lophelia Reef, Pinnacle #113 (depth 777 m, relief 61 m). (from Reed et al., 2004b)
146 ft tall Lophelia Mound 0.3 nm N-S
-2300 feet
-2500 feet
19
The second dive site (JSL II-3336) at Pinnacle #151 (28o17.0616’N, 79o36.8306’W) was also a deep-water Lophelia coral reef comprised entirely of coral and sediment (Table 1). Maximum depth was 758 m, with 44 m relief, and ~0.3 nm wide (N-S). The top was a series of ridged peaks from 713 to 722 m in depth. The lower flanks of the south face was a 10-20o slope of fine light colored sand with a series of 1-3 m high sand dunes or ridges that were linear NW-SE. The ridges had ~50% cover of thickets of Lophelia pertusa coral. The thickets consisted of 1 m tall dead, standing and intact, Lophelia pertusa colonies. Approximately 1-10% were alive on the outer parts (15-30 cm) on top of the standing dead bases. There was very little broken dead coral rubble in the sand and there was no evidence of trawl or mechanical damage. Most of the coral was intact, and the dead coral was brown. The sand between the ridges was fine and light colored, with 7-15 cm sand waves. The upper slope steepened to 45o and 70-80o slope near the upper 10 m from the top. The top of the pinnacle had up to 100% cover of 1-1.5 m tall coral thickets, on a narrow ridge that was 5-10 m wide. The coral consisted of both Lophelia pertusa and Enallopsammia profunda. Approximately 10-20% cover was live coral of 30-90 cm. The north slope was nearly vertical (70-80o) for the upper 10 m then consisted of a series of coral thickets on terraces or ridges. No exposed rock was visible and the entire pinnacle appeared to be a classic Lophelia mud mound.
No discernable zonation of macrobenthic fauna was apparent from the base to the top. Corals consisted of Lophelia pertusa, Enallopsammia profunda, Madrepora oculata, and some stylasterine hydrocorals. Dominant octocoral gorgonacea included Primnoidae (2 spp.), Isididae bamboo coral (Isidella sp. and Keratoisis flexibilis), and the alcyonaceans Anthomastus sp. and Nephthya sp (Table 3). Dominant sponges consisted of several species of Hexactinellida glass sponges, large yellow demosponges (60-90 cm diameter), Pachastrellidae, and Phakellia sp. fan sponges. Echinoderms included urchins (cidaroid and Hydrosoma? sp.) and comatulid crinoids, but no stalked crinoids. Some large decapod crustaceans included Chaceon fenneri and large galatheids. No mollusks were observed but were likely within the coral habitat that was not collected. Common fish were 2 m sharks, 25 cm eels, 25 cm skates, chimaera, and blackbelly rosefish (Table 4).
Region G: Miami Terrace Escarpment (from Reed et al., 2004b) The Miami Terrace is a 65-km long carbonate platform that lies between Boca Raton and South Miami at depths of 200-400 m in the northern Straits of Florida. It consists of high-relief Tertiary limestone ridges, scarps and slabs that provide extensive hard bottom habitat (Uchupi, 1966, 1969; Kofoed and Malloy, 1965; Uchupi and Emery, 1967; Malloy and Hurley, 1970; Ballard and Uchupi, 1971; Neumann and Ball, 1970). At the eastern edge of the Terrace, a high-relief, phosphoritic limestone escarpment of Miocene age with relief of up to 90 m at depths of 365 m is capped with Lophelia pertusa coral, stylasterine hydrocoral (Stylasteridae), bamboo coral (Isididae), and various sponges and octocorals (Reed et al., 2004b; Reed and Wright, 2004). Dense aggregations of 50-100 wreckfish were observed here by the PI during JSL submersible dives in May 2004 (Reed et al., 2004b). Previous studies in this region include geological studies on the Miami Terrace (Neumann and Ball, 1970; Ballard and Uchupi, 1971) and dredge- and trawl- based faunal surveys in the 1970s primarily by the University of Miami (e.g., Halpern, 1970; Holthuis, 1971, 1974; Cairns, 1979). Lophelia mounds are also present at the base of the escarpment (~670 m) within the axis of the Straits of Florida, but little is known of their
20 distribution, abundance or associated fauna. Using the Aluminaut submersible, Neumann and Ball (1970) found thickets of Lophelia, Enallopsammia (=Dendrophyllia), and Madepora growing on elongate depressions, sand ridges and mounds. Large quantities of L. pertusa and E. profunda have also been dredged from 738-761 m at 26 o22' to 24'N and 79 o35' to 37'W (Cairns, 1979).
Recent JSL submersible dives and fathometer transects by the PI at four sites (Reed Site #BU4, 6, 2, and 1b) indicated the outer rim of the Miami Terrace to consist of a double ridge with steep rocky escarpments (Table 1; Fig. 6; Reed and Wright, 2004; Reed et al., 2004b). At Miami Terrace Site #BU4, the narrow N-S trending east ridge was 279 m at the top and had a steep 95 m escarpment on the west face. The east and west faces of the ridges were 30-40o slopes with some near vertical sections consisting of dark brown phosphoritic rock pavement, boulders and outcrops. The crest of the east ridge was a narrow plateau ~10 m wide. At Site #BU6, the crest of the west ridge was 310 m and the base of the valley between the west and east ridges was 420 m. At Site #BU2, the echosounder transect showed a 13 m tall rounded mound at a depth of 636 m near the base of the terrace within the axis of the Straits of Florida. The profile indicated that it is likely a Lophelia mound. West of this feature the east face of the east ridge was a steep escarpment from 567 m to 412 m at the crest. The west ridge crested at 321 m. Total distance from the deep mound to the west ridge was 2.9 nm. Site #BU1b was the most southerly transect on the Miami Terrace. An E-W echosounder profile at this site indicated a double peaked east ridge cresting at 521 m, then a valley at 549 m, and the west ridge at 322 m. The east face of the west ridge consisted of a 155 m tall escarpment (Fig. 14).
Figure 14. Echosounder profile of Miami Terrace Escarpment, Site #BU1b, west ridge (depth 549 m at base, relief 155 m). (from Reed et al., 2004b)
-1200 feet
500-ft escarpment, east slope of Miami Terrace
-1800 feet
21 There were considerable differences among the sites in habitat and fauna; however, in general, the lower slopes of the ridges and the flat pavement on top of the terrace were relatively barren. However, the steep escarpments especially near the top of the ridges were rich in corals, octocorals, and sponges. Dominant sessile fauna consisted of the following Cnidaria: small (15- 30 cm) and large (60-90 cm) tall octocoral gorgonacea (Paramuricea spp., Placogorgia spp., Isididae bamboo coral); colonial scleractinia included scattered thickets of 30-60 cm tall Lophelia pertusa (varying from nearly 100% live to 100% dead), Madrepora oculata (40 cm), and Enallopsammia profunda; stylasterine hydrocorals (15-25 cm); and Antipatharia (30-60 cm tall) (Table 3). Diverse sponge populations of Hexactinellida and Demospongiae included: Heterotella sp., Spongosorites sp., Geodia sp., Vetulina sp., Leiodermatium sp., Petrosia sp., Raspailiidae, Choristida, Pachastrellidae, and Corallistidae. Other motile invertebrates included Asteroporpa sp. ophiuroids, Stylocidaris sp. urchins, Mollusca, Actiniaria, and Decapoda crustaceans (Chaceon fenneri and Galatheidae). Schools of ~50-100 wreckfish (Polyprion americanus), ~60-90 cm in length, were observed on several submersible dives along with blackbelly rosefish, skates, sharks, and dense schools of jacks (Table 4).
Region H: Pourtalès Terrace Lithoherms (from Reed et al., 2004a) The Pourtalès Terrace provides extensive, high-relief, hard-bottom habitat, covering 3,429 km2 (1,000 nm2) at depths of 200-450 m. The Terrace parallels the Florida Keys for 213 km and has a maximum width of 32 km (Jordan, 1954; Jordan and Stewart, 1961; Jordan et al., 1964; Gomberg, 1976; Land and Paull, 2000). Reed et al. (2004a) surveyed several deep-water, high-relief, hard- bottom sites including the Jordan and Marathon deep-water sinkholes on the outer edge of the Terrace, and five high-relief bioherms on its central eastern portion (Table 2, Fig. 7). The JSL and Clelia submersibles were used to characterize coral habitat and describe the fish and associated macrobenthic communities. These submersible dives were the first to enter and explore any of these features. The upper sinkhole rims range from 175 to 461 m in depth and have a maximum relief of 180 m. The Jordan Sinkhole may be one of the deepest and largest sinkholes known. The high-relief area of the middle and eastern portion of the Pourtalès Terrace is a 55 km-long, northeasterly trending band of what appears to be karst topography that consists of depressions flanked by well defined knolls and ridges with maximum elevation of 91 m above the terrace (Jordan et al., 1964; Land and Paull, 2000). Further to the northeast of this knoll-depression zone is another zone of 40-m high topographic relief that lacks any regular pattern (Gomberg, 1976). The high-relief bioherms (the proposed HAPC sites within this region) lie in 198 to 319 m, with a maximum height of 120 m. A total of 26 fish taxa were identified from the sinkhole and bioherm sites (Table 4). Species of potential commercial importance included tilefish, sharks, speckled hind, yellow-edge grouper, warsaw grouper, snowy grouper, blackbelly rosefish, red porgy, drum, scorpion fish, amberjack, and phycid hakes. Many different species of Cnidaria were recorded, including Antipatharia black corals, stylasterine hydrocorals, octocorals, and one colonial scleractinian (Solenosmilia variabilis) (Table 3).
Tennessee and Alligator Humps, Bioherms #1-4- Pourtalès Terrace (from Reed et al., 2004a) The Tennessee and Alligator Humps are among dozens of lithoherms that lie in a region called “The Humps” by local fishers, ~14 nm south of the Florida Keys and south of Tennessee and Alligator Reefs (Table 2, Fig. 7). Three dives were made by the PI on Bioherm #3 (Clelia 597, 598, 600; Aug. 2001), approximately 8.5 nm NE of Bioherm #2 (Fig. 15). Bioherm #3 consisted of two peaks 1.05 nm apart with a maximum relief of 62 m. The North Peak’s minimum depth
22 Figure 15. Echosounder profile of Pourtalès Terrace, Tennessee Bioherm #2 (depth 212 m at base, relief 85 m). (from Reed et al., 2004a)
was 155 m (submersible DGPS: 24o42.4573’N, 80o31.0513’W) and was 653 m wide at the base, which was 217 m deep at the east base and 183 m at the west side. The minimum depth of South Peak was 160 m and was about 678 m in width E to W at the base. The surrounding habitat adjacent to the mounds was flat sand with about 10% cover of rock pavement. From 213 m to the top, generally on the east flank of the mound, were a series of flat rock pavement terraces at depths of 210, 203, 198, 194, 183, and 171 m and the top plateau was at 165 m. Between each terrace a 30-45o slope consisted of either rock pavement or coarse sand and rubble. Below each terrace was a vertical scarp of 1-2 m where the sediment was eroded away leaving the edge of the terrace exposed as a horizontal, thin rock crust overhang of <1 m and 15-30 cm thick. The top of the bioherm was a broad plateau of rock pavement with 50-100% exposed rock, few ledges or outcrops, and coarse brown sand. Less time was spent on the western side, which was more exposed to the strong bottom currents. The west side of South Peak sloped more gradually than the eastern side, had more sediment, and no ledges were observed.
Fish Communities (from Reed et al., 2004a) A total of 31 fish taxa, of which 24 were identified to species level, were identified from our submersible videotapes and were associated with the deep-water sinkholes and high-relief bioherms (Table 4). Few studies have directly documented deep-water fish associations with deep-water reef habitats in the western Atlantic. Most of the work has concentrated on the Charleston Bump region of the Blake Plateau off Georgia and South Carolina (Sedberry, 2001). Ross (pers. comm.) reported the following species are common to both the deep-water Lophelia reefs on the Blake Plateau off the Carolinas and those of this study: Chloropthalmus agassizi, Helicolenus dactylopterus, Hoplostethus sp., Laemonema melanurum, Nezumia sp., and Xiphias gladius.
23 Species most common to the high-relief bioherms included deepbody boarfish, blueline tilefish, snowy grouper, and roughtongue bass. Some species were common at both the sinkhole and bioherm sites and included snowy grouper, blackbelly rosefish, and mora. In addition to the moribund swordfish observed in the Jordan Sinkhole, a swordfish was observed from the NR-1 submersible on top of Pourtales Terrace (C. Paull, pers. observation).
Species of potential commercial importance included tilefish, sharks, speckled hind, yellowedge grouper, warsaw grouper, snowy grouper, blackbelly rosefish, red porgy, drum, scorpionfish, amberjack, and phycid hakes. However, the fish densities that we saw at any of the sites were in insufficient numbers to suggest commercial or recreation harvest. In fact, any of the features, both sinkholes and bioherms, could be overfished very easily since only a few individuals of the larger grouper species were present at any one site.
Benthic Communities (from Reed et al., 2004a) The benthos at the bioherm sites was dominated by sponges, octocorals and stylasterids (Table 3). A total of 21 taxa of Cnidaria were sampled or observed and 16 were identified to species level. These included 3 species of antipatharian black coral, 5 stylasterid hydrocorals, 11 octocorals with one possible new species, and 1 scleractinian (Solenosmilia variabilis). Eight species were associated only with the Pourtalès sinkholes and not the bioherms; these included two species of antipatharians; the octocorals Paramuricea placomus, Plumarella pourtalesii, Trachimuricea hirta; and the scleractinian Solenosmilia variabilis. Although Gomberg (1976) found evidence of skeletal remains of the colonial scleractinians Lophelia and Madrepora in sediment samples from the terrace, we did not see any colonies at our dive sites. Sponges identified from collections included 28 taxa. Five species of stylasterine hydrocorals were Distichopora foliacea, Pliobothrus echinatus, Stylaster erubescens, S. filogranus, and S. miniatus. On the flat pavement adjacent to the base of the mounds, stylasterids and antipatharian black coral bushes were common along with sea urchins and sea stars.
The densities of sponges, stylasterid hydrocorals and octocorals were very high, especially on the plateaus and terraces of the bioherms on the Pourtalès Terrace. Maximum densities of sponges (>5 cm) on the plateaus ranged from 1-80 colonies m-2. Stylasterid coral densities ranged from 9-96 colonies m-2 and octocorals 16-48. Densities of sponges (1-2 colonies m-2) and stylasterids (1-20) also dominated the terraces and slopes of the bioherm sites but generally in lower densities than the peak plateaus whereas the octocorals generally had higher densities on the flanks (1-80 colonies m-2).
Region I: Southwest Florida Shelf Lophelia Lithoherms (from Reed et al., 2003; Reed et al., 2004 a, b, d) This region consists of dozens and possibly hundreds of 5-15 m tall lithoherms at depths of 500 m, some of which are capped with thickets of live and dead Lophelia coral (Fig. 8). In 1987, Newton et al. described the area from limited dredge and seismic survey. In 2003, Seabeam topographic mapping was conducted by the PI over a small portion of the region (Table 1, Figs. 16,17); ROV dives ground-truthed three of the features: a 36-m tall escarpment and two of the lithoherms (Reed, 2004; Reed et al., 2003; Reed et al., 2004b,d). The lithoherms appeared to consist of rugged black phosphorite-coated limestone boulders and outcrops capped with 0.5-1.0 m tall thickets of Lophelia pertusa, which were up to ~10-20% live. Dominant sessile
24 macrofauna included stony corals, octocorals, stylasterid hydrocorals, black corals and sponges (Table 3). The high number of hard bottom lithoherms revealed by the (limited) Seabeam mapping effort indicated tremendous potential for unexplored coral and fish habitat in this region.
Figure 16. Seabeam image of escarpment and lithoherms at Region I off southwest Florida slope. ?= Innovator ROV dive sites #6- 8. (from Reed et al., 2004b)
Figure 17. Seabeam image of escarpment and lithoherms at Region I off southwest Florida slope; simulated view from top of escarpment, looking south.. (from Reed et al., 2004b)
Lophelia Lithoherms
100 ft tall Escarpment
25 An ROV dive by the PI on the 36-m tall escarpment (Fig. 17; top- 412 m, base- 448 m), showed a near vertical wall with a series of narrow ledges, and very rugged topography with crevices and outcrops. Dominant sessile fauna consisted of Antipatharia black coral (30 cm tall), numerous octocoral gorgonacea including Isididae bamboo coral (30-40 cm), and sponges (Heterotella sp., Phakellia sp., Corallistidae). Pinnacle #4 was a 12 m tall and 60 m wide lithoherm at a depth of 466 m. Eight other lithoherms were apparent on the ROV’s sonar within a 100 m radius. A transect up the face of the pinnacle revealed a series of terraces on a rugged 45o up to 70o rock slope which consisted of black rock boulders (1-2 m) and outcrops with 1 m crevices. The top ridge was oriented ~NNE. Thickets of live and dead Lophelia pertusa were found on some of the slope terraces but primarily on the top ridge. The NE slope face appeared to have more live coral than the NW face. Some of the thickets were ~30-60 cm tall and 60-90 cm diameter. Coral cover was estimated from <5% to over 50% in some areas, and estimated to be 1-20% live. The dominant fauna were similar to the escarpment except for Lophelia which was not observed on the escarpment. Common sessile benthic species included Cnidaria: Antipatharia black coral (Antipathes sp. and Cirrhipathes sp.), Lophelia pertusa, gorgonacea octocorals; and sponges: Heterotella sp. and other Hexactinellida vase sponges, various plate and vase Demospongiae (Pachastrellidae, Petrosiidae, Choristida). Common motile invertebrates included Mollusca, Holothuroidea, Crinoidea, Decapoda crustaceans (Chaceon fenneri and Galatheidae), blackbelly rosefish, and various other benthic fish (fish tapes have not been analyzed yet).
SUMMARY AND RECOMMENDATIONS
The biological and geological characteristics of six regions of deep-water reefs off the southeastern U.S.A. from southwest Florida to South Carolina were summarized in this report based on current data and knowledge compiled primarily from recent submersible and ROV dives. Region A, the Oculina Reefs, have been designated an Habitat Area of Particular Concern since 1984 (NOAA, 1982; Reed, 1981d; Reed, 2002b) and portions are a Marine Protected Area for the protection of the coral habitat and snapper/ grouper complex. Even so, extensive areas of the Oculina reefs have been severely impacted by legal and illegal bottom trawling since 1984. The six regions outlined in this report (Regions B-D, G-I) are each unique in their own respect. The resource potential of the deep-water habitats in this region is unknown in terms of fisheries and novel compounds yet to be discovered from associated fauna that may be developed as pharmaceutical drugs. Although these habitats are not currently designated as MPAs or HAPCs, they are incredibly diverse and irreplaceable resources. Activities involving bottom trawling, pipelines, or oil/gas production could negatively impact these reefs. This PI strongly recommends that HAPC designation be given to these deep-water reef habitats to provide some protection to these resources. Evidence of potential spawning aggregations of wreckfish (Polyprion americanus) and considerable populations of blackbelly rosefish (Helicolenus dactylopterus) and other commercially important species could actually threaten the future longevity of these fragile habitats unless bottom trawling in these regions is prohibited or strictly regulated and monitored. These studies summarized in this report are only preliminary and point to the need for additional geological, biological and ecological research. Initially, most of these regions need detailed mapping and habitat characterization studies which will provide data for final determinations of potential HAPC boundaries and future research needs.
26 ACKNOWLEDGMENTS
Numerous individuals have contributed to this research over many years. I especially thank Dr. Robert Avent who initiated these studies in the 1970s and Dr. Charles 'Skip' Hoskin who provided years of enthusiastic collaboration and leadership. I gratefully acknowledge the Division of Biomedical Marine Research at Harbor Branch Oceanographic Institution (HBOI) which funded and provided data from recent JSL submersible dives on the Oculina and Lophelia reefs and lithoherms. We thank NOAA’s Office of Ocean Exploration for funding our biomedical research studies on these reefs in 2002 and 2003 and the State of Florida for funding the Center of Excellence in Biomedical and Marine Biotechnology (HBOI and Florida Atlantic University) which provided ship and submersible funding in 2004 and 2005. The following individuals provided taxonomic identifications: Dr. Shirley Pomponi and Dr. Michelle Kelly, sponges; Dr. Charles G. Messing and John Miller, echinoderms; Dr. Stephen Cairns, scleractinia; and Dr. Charles G. Messing, gorgonacea. The various crew of HBOI vessels, and Johnson-Sea- Link and Clelia submersibles are also thanked for their support. This is contribution no. 1570 from Harbor Branch Oceanographic Institution.
27 Table 1. Site summary for deep-water coral reefs and lithoherms off SE USA. In order north to south. Site #1-33 refer to Fig. 4. (from Reed, 2002a; Reed et al., 2004a,b)
*Site Depth at Depth at Max. Relief GPS Coordinates Reference Base Peak (m); (Width (Peak) (m) (m) at base) Region D o 1) Stetson’s Reefs, Stetson’s 780 627 153 32 01.6882’N, o Pinnacle (0.8 nm N-S) 77 39.6648’W
2) Stetson’s Reefs, Pinnacle 694 579 114 32o00.6302’N, #3, Peak 1-4 (Peak 1) (2.2 nm N-S) 77o41.9285’W (Peak 1)
Region C o 3) Savannah Lithoherms, 550 500 54 31 48’N, o ALVIN site 79 15’W
4) Savannah Lithoherms, Site 549 511 38 31o44.3814’N, 2, Pinnacle #6 (0.4 nm NE- 79o05.2516’W SW)
5) Savannah Lithoherms, 549 533 15 31o44.0975’N, Site 2, Pinnacle #5 (0.3 nm NE- 79o05.5544’W SW)
6) Savannah Lithoherms, Site 537 487 50 31o42.2555’N, 2, Pinnacle #1 (0.53 nm N-S) 79o07.4831’W
7) Savannah Lithherms 541 31o41.82’N, 79o08.60’W
8) Savannah Lithoherms 532 499 33 31o41.5’N, 79o18.06’W
9) Savannah Lithoherms, Site 549 488 61 31o41.4259’N, 1, Pinnacle #4 (0.47 nm N-S) 79o08.5964’W
10) Savannah Lithoherms 503 490 13 31o41.23’N, 79o17.46’W
Region B o 11) Paull (2000) Lithoherm 671 579 91 30 48.2’N o Site (440-914) (150 max) 79 38.4’W
12) Jacksonville Lophelia 701 544 157 max; Peak 30o30.1194’N, Reef, Pinnacle #204B, Peak 6 6= 107 79o39.4743’W (3nm N-S; 0.8nm E-W)
13) Jacksonville Lophelia 866 744 122 (0.9 nm 30o16.8114’N, Reef, Pinnacle #186 N-S; 0.9 nm E- 79o38.9784’W W)
28
14) St. Augustine Lophelia 822 734 88 29o40.2628’N, Reef, Pinnacle #3 (0.99 nm N-S) 79o38.0678’W
15) Cape Canaveral Lophelia 777 716 61 28o47.6258’N, Reefs, Pinnacle #113 (0.3 nm N-S; 0.9 79o37.5859’W nm NW-SE)
16) Cape Canaveral Lophelia 793 762 30 28o46.72’N, Reefs 79o41.17’W
17) Cape Canaveral Lophelia 791 716 75 (0.53 nm 28o39.8464’N, Reef, Pinnacle #129 N-S) 79o37.6735’W
18) Cape Canaveral Lophelia 762 718 44 (0.78 nm 28o28.3513’N, Reef, Pinnacle #TS7 (Near P N-S) 79o37.0064’W 135)
19) Cape Canaveral Lophelia 758 713 44 28o17.0616’N, Reefs, Pinnacle #151 (0.3 nm N-S) 79o36.8306’W
20) Cape Canaveral Lophelia 838 741 97 28o02.04’N, Reefs 79o36.51’W (Loran C)
21) Ft. Pierce Lophelia Reef, 750 721 29 27o39.4305’N, Pinnacle #TS4 (near P212) (0.84 nm N-S) 79o34.9679’W
22) Stuart Lophelia Reef, 723 676 46 (0.95 nm 27o12.5695’N, Pinnacle #292 N-S; 0.82 nm 79o35.5994’W E-W)
23) Jupiter Lophelia Reef, 723 685 42 (1.66 nm 27o01.3474’N, Pinnacle #293 N-S; 1.0 nm E- 79o35.3889’W W)
Region A o o Oculina Reefs 70-100 24 27 32.8’N, 79 56.2’W to o o (Reed, 1980, 2002a,b) 28 59.2’N, 80 06.6’W
Region E o o (Mullins et al., 1981 ; Reed, 1000- 40 27 40’N, 78 15’W to o o 2002a) 1300 27 10’N, 77 30’W
Region F o o (Neumann et al., 1977; 610- 675 50 26 56.72’N, 79 16.02’W to o o Messing et al., 1990; Reed, 27 25’N, 79 20’W 2002a)
Region G o 24) Miami Terrace, East 375 279 95 26 05.7066’N, o Ridge, W. Face, Site #BU4 79 50.3634’W (ridge top)
29
25) Miami Terrace, East 335 284 51 26o05.6902’N, Ridge, E. Face, Site #BU4 79o50.2540’W (base of escarptment)
26) Miami Terrace, West 437 310 126 26o01.2885’N, Ridge, East Face, Site #BU6 79o49.3258’W (base of escarpment)
27) Miami Terrace, East 573 399 174 25o41.9970’N, Ridge, E. Face, Site #BU2 79o51.0510’W (base of escarpment)
28) Miami Terrace, West 391 321 70 25o41.9959’N, Ridge, E. Face, Site #BU2 79o51.8924’W (base of escarpment)
29) Miami Terrace, West 549 393 155 25o35.9963’N, Ridge, Base E. Face, Site 79o52.9368’W (base of BU1b escarpment)
30) Miami Terrace, West 430 322 112 25o35.9864’N, Ridge, W. Face, Site #BU1b 79o54.2491’W
Region H o o *Pourtales Terrace Sites 198- 461 12- 24 15.33’N, 80 54.27’W to o o (Reed et al., 2004) 180 24 44.71’N, 80 27.59’W
Region I o 31) SW Fla. Lithoherms, 558 554 4 26 19.9094’N, o Pinnacle #1 84 45.8639’W
32) SW Fla. Lithoherms, Site 448 412 36 escarp- 26o20.3915’N, 2 Escarpment ment 84o44.8733’W
33) SW Fla. Lithoherms, 466 454 12 26o20.0133’N, Pinnacle #4 84o45.0030’W (base)
Regions A-H: Southeast USA; Region I: Eastern Gulf of Mexico; *= Region I, Pourtales Terrace Sites- see separate table; dive number: JSL I, II= HBOI’s Johnson-Sea-Link I and II manned submersibles, CORD= HBOI’s Cord Remotely Operated Vehicle (ROV), ROV= Sonsub Innovator ROV, ALVIN= WHOI’s Alvin submersible; depth= at base, peak, maximum relief, and maximum width at base of bioherm; coordinates are submersible/ROV GPS location at peak of bioherm (or as indictated).
30 Table 2. Site summary for deep-water sinkholes and bioherms off south Florida, Pourtalès Terrace. (from Reed et al., 2004a) *Site Reference Depth Max. Width GPS Coordinates (m) Relief (m) (m) Naples Sinkhole 175 -55 152 26o05.1791’N 84o13.4678’W
Jordan Sinkhole 366 -180 229 24o16.4241’N, 81o02.1846’W
Marathon Sinkhole 461 -61 610 24o15.3289’N, 80o54.2705’W
Key West Bioherm 198 12 422 24o21.8038’N, 81o50.7397’W
Tennessee Bioherm #1 319 120 574 24o30.1670’N, 80o40.1880’W
Tennessee Bioherm #2 213 85 1613 24o35.2676’N, 80o35.3345’W
Alligator Bioherm #3 217 62 678 24o42.4573’N, 80o31.0513’W
Alligator Bioherm #4 213 48 1778 24o44.71’N, 80o27.59’W
Depth and width at base of bioherm or top of sinkhole; coordinates are submersible GPS location at peak of bioherm or base of sinkhole.
31 Table 3. Species list of macroinvertebrates associated with deep-water reefs off southeastern U.S.A. (Phyla: ART= Arthropoda, BRY= Bryozoa, CNI= Cnidaria, ECH= Echinodermata, MOL= Mollusca, POR= Porifera, VES= Vestimetifera; Sites: SC= Stetson’s Reefs, South Carolina; GA= Savannah Lithoherms, Georgia; FL-E= East Coast Florida Lophelia Reefs; MT= Miami Terrace Escarpment; PT= Pourtalès Terrace Sinkholes and Bioherms; FL-W= SW Florida Lithoherms; VK= Viosca Knoll). (from Reed et al., 2004a,b) Phylum Taxonomy Min Max SC GA FL-E MT PT FL-W VK Depth Depth (m) (m) ART Chaceon fenneri (golden crab) 509 509 X BRY Membranipora? sp. Blainville, 1830 631 631 X CNI Muriceides sp. (not hirta, not kukenthali) Studer, 1887 191 191 X CNI Stylaster erubescens Pourtales, 1868 175 186 X CNI Swiftia casta (Verrill, 1883) 525 525 X CNI Swiftia new sp.? Duchassaing & Michelotti, 1864 497 497 X CNI Solenosmilia variabilis Duncan, 1873 470 470 X CNI Trachymuricea hirta (Pourtales, 1867) 462 468 X CNI Paramuricea placomus (Linnaeus, 1924) 462 470 X CNI Antipathes rigida? Pourtales, 1868 319 319 X CNI Placogorgia mirabilis Deichmann, 1936 172 212 X CNI Thesea parviflora Deichmann, 1936 183 183 X CNI Hydroida 202 656 X X CNI Stylaster miniatus (Pourtales, 1868) 175 200 X CNI Stylaster filogranus Pourtales, 1871 175 200 X CNI Distichopora foliacea Pourtales, 1868 175 175 X CNI Pliobrothus echinatus Cairns, 1986 175 175 X CNI Bathypsammia? sp. Marenzeller, 1907 418 640 X X CNI Clavularia new sp.? Quoy & Gaimard, 1834 648 648 X CNI Eunephthya nigra (Pourtales, 1868) 648 768 X CNI Octocorallia, unid. spp. 501 671 X X CNI Lophelia pertusa (Linnaeus, 1758) 284 815 X X X X X X CNI Scleractinia, unid. spp. 582 632 X X CNI Enallopsammia profunda (Pourtales, 1867) 305 742 X X X X CNI Ifalukellidae, new sp.? Bayer, 1955 (ye morph) 502 649 X X CNI Eunicella modesta (Verrill, 1883) 518 732 X X CNI Keratoisis flexibilis (Pourtales, 1868) (wh morph) 378 816 X X X X CNI Ifalukellidae, new sp.? Bayer, 1955 (or morph) 519 656 X X CNI Actiniaria 565 751 X CNI Placogorgia? sp.1 Wright & Studer, 1889 565 579 X CNI Chrysogorgia squamata (Verrill, 1883) 581 581 X CNI Bathypathes alternata Brook, 1889 466 716 X X CNI Pterostenella? new sp.? Versluys, 1906 754 754 X CNI Zoanthidea, unid. sp.2 734 734 X CNI Stylaster unid. sp.1 557 557 X CNI Placogorgia tenuis? (Verrill, 1883) 457 557 X CNI Callogorgia verticillata (Pallas) 511 511 X CNI Isidella sp.1 Gray, 1857 744 762 X CNI Paramuricea sp.2 Kölliker, 1865 573 573 X
32 CNI Madrepora oculata Linnaeus, 1758 322 763 X X X CNI Paramuricea sp.4 Kölliker, 1865 762 762 X CNI Plumarella pourtalessi (Verrill, 1883) 171 753 X X X X X CNI Keratoisis flexibilis (Pourtales, 1868) (pi morph) 374 734 X X CNI Actiniaria, unid. sp.1 (Venus fly trap) 284 734 X X CNI Candidella imbricata (Johnson, 1862) + Thouarella? sp. 732 732 X Gray, 1870 CNI Paramuricea sp.3 Kölliker, 1865 558 732 X CNI Anthomastus nr. agassizzi Verrill, 1922 420 753 X X CNI Telestula? sp.2 Madsen, 1944 734 784 X CNI Paramuricea sp.5 Kölliker, 1865 743 744 X CNI Paramuricea sp.1 Kölliker, 1865 590 744 X CNI Paramuricea sp.6 Kölliker, 1865 328 727 X X CNI Paramuricea sp.7 Kölliker, 1865 711 711 X CNI Paramuricea sp.8 Kölliker, 1865 701 716 X CNI Capnella nigra (Pourtales, 1868) 325 762 X X CNI Paramuricea multispina Deichmann, 1936 189 715 X X CNI Plexauridae, unid. sp.1 Gray, 1859 579 716 X X CNI Muriceides hirta? (=Trachymuricea) (Pourtales, 1867) 681 716 X CNI Paramuriceidae sp.2 (nr. Paramuricea echinata 716 716 X Deichmann, 1936) CNI Paramuriceidae sp.4 (nr. Paramuricea placomus 296 296 X (Linnaeus)) CNI Antipatharia, unid. sp.1 (re-or morph) 283 767 X X X X X CNI Paramuriceidae sp.3 (nr. Paramuricea placomus 283 304 X (Linnaeus)) CNI Antipatharia, unid. sp.2 (wh-pi morph) 328 515 X X X X CNI Paramuriceidae sp.5 (nr. Echinomuricea atlantica 284 284 X (Johnson, 1862)) CNI Zoanthidea, unid. sp.1 419 699 X X CNI Paramuricea sp.9 Kölliker, 1865 326 336 X CNI Paramuriceidae sp.6 (nr. Paramuricea placomus 326 326 X (Linnaeus)) CNI Paramuriceidae sp.7 (nr. Paramuricea multispina 323 323 X Deichmann, 1936) CNI Zoanthidea, unid. sp.3 328 328 X CNI Villogorgia nr. nigrescens Duchassaing & Michelotti, 1860 215 215 X CNI Paramuricea sp.10 Kölliker, 1865 403 403 X CNI Paramuricea sp.11 Kölliker, 1865 322 358 X CNI Paramuricea sp.12 Kölliker, 1865 366 366 X CNI Stylasteridae, unid. sp. 173 742 X X X X CNI Paramuriceidae sp.8 (nr. Echinomuricea atlantica 323 323 X (Johnson, 1862)) CNI Paramuricea sp.13 Kölliker, 1865 323 323 X CNI Hydroida, unid. sp.1 284 322 X ECH Holothuroidea 181 181 X ECH Tamaria? sp. 653 653 X ECH Solaster sp. 653 653 X ECH Asteroidea + Cidaroidea 516 516 X ECH Asteroidea, 2 unid. spp. 518 518 X ECH Asteroidea, unid. sp.1 454 454 X ECH Asteroporpa? sp. 304 304 X
33 MOL Calliostoma pulchrum (C.B. Adams, 1850) 187 187 X MOL Hyalina albolineata (Orbigny, 1842) 187 187 X MOL Scaphella gouldiana (Dall, 1887) 187 188 X MOL Bivalvia, unid. sp.1 445 445 X MOL Bursa tenuisculpta (Dautzenberg & Fischer, 1906) 187 283 X X MOL Perotrochus amabilis (F.M. Bayer, 1963) 181 265 X MOL Conus villepini Fisher and Bernardi, 1857 171 188 X MOL Murex beauii Fischer & Bernardi, 1857 188 188 X MOL Entemnotrochus adansonianus (Crosse & Fischer, 1861) 180 265 X MOL Perotrochus midas F.M. Bayer, 1965 262 393 X POR Haplosclerida? 171 184 X POR Aka sp. de Laubenfels, 1934 or Spongosorites sp. 543 543 X Topsent, 1896 + Haplosclerida POR Haplosclerida + Siphonodictyon sp. Bergquist, 1965 or 187 543 X Spongosorites sp. Topsent, 1896 POR Theonellidae 470 472 X POR Pachastrella sp. Schmidt, 1868 or Poecillastra sp. Sollas, 467 467 X 1888 POR Stellettidae? 312 312 X POR Erylus transiens (Weltner, 1882) 262 262 X POR Halichondrida 260 260 X POR Theonellidae, new genus, new sp. 199 208 X POR Mycalidae 284 312 X POR Chondrosia? sp. Nardo, 1847 297 300 X POR Halichondriidae 237 648 X X POR Plakortis sp. Schulze, 1880 220 312 X POR Petrosiidae 178 750 X X X X X POR Porifera, unid. sp. 192 297 X POR Corallistes sp. Schmidt, 1870 or Callipelta sp. Sollas, 1888 206 206 X POR Spirophorida 183 183 X POR Lithistida 185 310 X POR Geodiidae 180 816 X X POR Poecilosclerida 132 717 X X POR Epipolasis sp. de Laubenfels, 1936 211 211 X POR Axinellida + Plakortis? sp. Schulze, 1880 210 210 X POR Axinellidae 168 183 X POR Characella? sp. Sollas, 1886 198 198 X POR Stellettinopsis? sp. Carter, 1879 198 198 X POR Echinodictyum sp. Ridley, 1881 171 172 X POR Phakellia new sp.1 Bowerbank, 1862 171 171 X POR Auletta sp. Schmidt, 1870 171 207 X POR Phakellia new sp.2 Bowerbank, 1862 174 174 X POR Phakellia new sp.3 Bowerbank, 1862 174 174 X POR Dictyoceratida? 172 172 X POR Pachastrellidae 166 811 X X X X X X POR Lychniscosida 649 662 X POR Lyssacinosida 628 757 X X POR Phakellia sp. Bowerbank, 1862 171 756 X X X X X X POR Corallistes sp. Schmidt, 1870 226 689 X X POR Oceanapia sp. Norman, 1869 172 652 X X
34 POR Plakinidae 638 660 X POR Aka (Siphonodictyon) sp.de Laubenfels, 1934 183 648 X X POR Ancorina? sp. Schmidt, 1862 641 641 X POR Phakellia sp.2 Bowerbank, 1862 509 509 X POR Hexasterophora 517 761 X X X POR Axinellida 201 499 X X POR Biemnidae 512 628 X X POR Pachastrellidae (different) 527 527 X POR Ircinia new sp.? Nardo, 1833 500 500 X POR Choristida, new sp.? 520 520 X POR Raspailiidae 321 763 X X X X POR Hexactinellida 186 800 X X X X X POR Heterotella sp. Gray, 1867 418 762 X X X POR Stylocordyla sp. Thomson, 1873 515 515 X POR Phakellia sp.3 Bowerbank, 1862 515 515 X POR Aka sp. de Laubenfels, 1934 + Hadromerida 456 456 X POR Myxillina? sp. Hajdu, Van Soest & Hooper, 1994 442 442 X POR Dendroceratida 448 448 X POR Hyalonematidae? + Zoanthidea 737 737 X POR Oceanapiidae 758 758 X POR Calthropellidae 757 757 X POR Ancorinidae? 586 586 X POR Dercitus cf. bucklandi (Bowerbank, 1858) 809 809 X POR Aphrocallistes sp. Gray, 1858 587 800 X POR Polymastia sp. Bowerbank, 1864 726 726 X POR Phakellia sp. (different) Bowerbank, 1862 735 735 X POR Corallistidae 186 767 X X X POR Asterophorida 431 431 X POR Leiodermatium sp. Schmidt, 1870 172 754 X X X POR Spongosorites sp. Topsent, 1896 171 671 X X X X POR Geodia sp. Lamarck, 1815 174 767 X X X POR Hexactinellida + Zoanthidea 328 411 X POR Poecillastra? sp. Sollas, 1888 323 427 X X POR Choristida 173 509 X X X POR Choristidae? 323 323 X POR Oceanapiidae or Topsentia sp. Berg, 1899 173 173 X POR Hymedesmia sp.1 Bowerbank, 1864 (blue morph) 172 179 X POR Hymedesmia sp.2 Bowerbank, 1864 (ye morph) 172 179 X POR Demospongiae 170 541 X X POR Discodermia sp. du Bacage, 1869 180 269 X POR Choristida or Petrosida 258 258 X POR Zyzzya sp. de Laubenfels, 1936 222 222 X POR Smenospongia sp. Wiedenmayer, 1977 or Ircinia sp. 222 222 X Nardo 1833 POR Petrosida or Halichondrida 183 183 X POR Vetulina sp. Schmidt, 1879 or Leiodermatium sp. Schmidt, 415 415 X 1870 POR Erylus sp. Gray, 1867 216 356 X X VES Vestimentifera, unid. sp. 443 443 X
35 Table 4. Species list of fish associated with deep-water reefs off Florida (Sites: FL= Florida East Coast Lophelia Reefs; MT= Miami Terrace Escarpment; PT= Pourtalès Terrace). (from Reed et al., 2004a,b)
Taxonomy Common Max Min FL MT PT Name Depth (m) Depth (m) Anthias nicholsi Firth, 1933 yellowfin bass 283 179 X X Antigonia capros Lowe, 1843 deepbody boarfish 219 174 X Beryx dacadactylus? alphonsino? 287 X Brotulidae cusk-eel 469 322 X X Carcharhinus falciformis (Müller & Henle, 1839) silky shark 522 335 X Caulolatilus microps Goode and Bean, 1878 blueline tilefish 223 172 X Chaetodon aya bank butterflyfish 179 X Chlorophthalmidae greeneye 296 X Chlorophthalmus agassizi Bonaparte, 1840 shortnose greeneye 522 396 X X Conger conger? conger eel 296 0 X Congridae conger eel 381 0 X Cookeolus japonicus (Cuvier, 1829) longfinned bulleye 198 171 X Epinephelus drummondhayi Goode and Bean, 1878 speckled hind 183 Epinephelus flavolimbatus Poey, 1865 yellowedge grouper 174 Epinephelus nigritus (Holbrook, 1855) Warsaw grouper 198 180 X Epinephelus niveatus (Valenciennes, 1828) snowy grouper 308 174 X Epinephelus sp. (misty grouper?) misty grouper? 287 X Galeus arae (Nichols, 1927) roughtail catshark 518 X Gephyroberyx darwinii (Johnson, 1866) big roughy 518 392 X Gymnothorax sp. (cf. funebris Ranzani, 1840) green moray 187 174 Gymnothorax sp. (new moray?) new moray 179 X Helicolenus dactylopterus (Delaroche, 1809) blackbelly rosefish 497 179 X X Hemanthias sp. seabass 194 174 X Hemanthias vivanus (Jordan & Swain, 1885) red barbier 191 168 X Hoplostethus mediterraneus Cuvier, 1829 silver roughy 461 X Hoplostethus sp. roughies 496 189 X Hydrolagus sp. spotted ratfish 762 714 X Hyperoglyphe sp. barrelfish 287 284 X Laemonema melanurum Goode and Bean, 1896 mora 546 186 X X X Mola mola ocean sunfish 180 X Mustelidae? dogfish 586 X Mustelus sp. dogfish 369 X Myctophidae laternfish 500 296 X X Nezumia sp. (3 spp.- N. bairdii, N. aequalis, or N. grenadier, rattail 726 322 X X X atlantica) Ostichthys trachypoma (Günther, 1859) bigeye soldierfish 180 Pagrus pagrus (Linnaeus, 1758) red porgy 175 Pareques iwamotoi Miller and Woods, 1988 blackbar drum 183 Peristidion sp. armored sea robin 438 X Plectranthias garrupellus Robins and Starck, 1961 apricot bass 172 X Polyprion americanus wreckfish 693 283 X X Pronotogrammus martinicensis (Guichenot, 1868) roughtongue bass 212 168 X
36 Raja s p. skate 738 339 X X Scorpaenidae scorpionfish 296 186 X X Scyliorhinidae? catshark? 326 X Seriola dumerili (Risso, 1810) greater amberjack 187 175 X Seriola rivoliana Almaco jack 179 X Squalidae dogfish 399 322 X Synaphobranchidae? cutthroat eel 762 714 X Unid.- silver body, barbels 336 X Urophycis sp. phycid hake 297 X Xeiidae? red dory? 376 X Xiphias gladius Linnaeus, 1758 swordfish 518 X
FIGURE CAPTIONS
Figure 1. Coral colony and branch tip: top- Oculina varicosa (80m); middle- Lophelia pertusa (490m); bottom- Enallopsammia profunda (585m). (scale lines = 1 cm; top left fig. Scale = 5 cm) (from Reed, 2002a; Hydrobiologia 471: 57-69)
Figure 2. Depth range and maximum relief of deep-water coral reefs off southeastern U.S.A. Dominant colonial coral listed for each site (see Figure 3 for site locations). (from Reed, 2002a; Hydrobiologia 471: 57-69) Figure 3. Deep-water coral reef regions off southeastern U.S.A. (see Table 1 for locations). ?= Johnson-Sea-Link I and II submersible dive sites; Regions: A=Oculina Coral Reefs, B= East Florida Lophelia Reefs, C= Savannah Lophelia Lithoherms, D= Stetson’s Reefs (D1= region of dense pinnacles), E= Enallopsammia Reefs (Mullins et al., 1981), F= Bahama Lithoherms (Neumann et al., 1977), G= Miami Terrace Escarpment. (from Reed et al., 2004a; chart from NOAA, NOS, 1986)
Figure 4. Submersible dive sites and echosounder sites on deep-water reefs off southeastern U.S.A. (see Table 1 for locations). ?# = Johnson-Sea-Link I and II submersible dive sites, F# = high-relief pinnacles from echosounder transect. (from Reed et al., 2004a; chart from NOAA, NOS, 1986)
Figure 5. Detailed chart of high-relief region with Lophelia coral mounds on Charleston Bump, Blake Plateau (from Popenoe and Manheim, 2001; American Fisheries Society Symposium 25: 43- 94)
Figure 6. Bathymetry and submersible dive sites on Miami Terrace Escarpment at Region G. (see Table 1 for locations). ?= Johnson-Sea-Link I submersible dive sites. (from Reed et al., 2004a; chart from Ballard and Uchupi, 1971; MTS Journal 5: 43-48)
Figure 7. Bathymetry and submersible dive sites on Pourtalès Terrace at Region H. (see Table 2 for locations). ?= Johnson-Sea-Link and Clelia submersible dive sites; JS= Jordan Sinkhole, MS= Marathon Sinkhole, T1= Tennessee Humps Bioherm #1, T2= Tennessee Humps Bioherm #2, A3= Alligator Humps Bioherm #3, A4= Alligator Humps Bioherm #4. (from Reed et al., 2004b; chart from Malloy and Hurley, 1970; Geol. Soc. Amer. Bull. 81: 1947-1972)
37
Figure 8. Deep-water coral lithoherms and ROV dive sites at Region I off southwest Florida slope (see Table 1 for locations). ?= Innovator ROV dive sites. (from Reed et al., 2004a; chart from NOAA, NOS, 1986)
Figure 9. Echosounder profile of Stetson’s Pinnacle (depth 780 m, relief 153 m). (from Reed et al., 2004b)
Figure 10. Echosounder profile of Savannah Lithoherm, Pinnacle #1 (depth 537 m, relief 50 m). (from Reed et al., 2004b)
Figure 11. Height of Lophelia pinnacles and lithoherms on echosounder transects from Jacksonville to Jupiter, Florida at depths of 600 to 800 m. (from Reed et al., 2004b)
Figure 12. Echosounder profile of Jacksonville Lithoherm, Pinnacle #204B (depth 701 m, relief 157 m). (from Reed et al., 2004a)
Figure 13. Echosounder profile of Cape Canaveral Lophelia Reef, Pinnacle #113 (depth 777 m, relief 61 m). (from Reed et al., 2004a)
Figure 14. Echosounder profile of Miami Terrace Escarpment, Site #BU1b, west ridge (depth 549 m at base, relief 155 m). (from Reed et al., 2004a)
Figure 15. Echosounder profile of Pourtalès Terrace, Tennessee Bioherm #2 (depth 213 m at base, relief 85 m). (from Reed et al., 2004b)
Figure 16. Seabeam image of escarpment and lithoherms at Region I off southwest Florida slope. ?= Innovator ROV dive sites #6 and 7. (from Reed et al., 2004b)
Figure 17. Seabeam image of escarpment and lithoherms at Region I off southwest Florida slope, simulated view from top of escarpment. ?= Innovator ROV dive sites #6 and 7. (from Reed et al., 2004b)
38 REFERENCES
Deep-Water Reefs - Habitat, Biological, and Geological References Part 1: Western Atlantic- North Carolina to Florida Part 2: Gulf of Mexico (p.64) Part 3: Eastern Atlantic and General Deep Sea Reefs (p.68)
Compiled by John Reed, October 20, 2004 [*- Reference SEAMAP Deep-water Florida Data Set]
PART I- Southeastern USA, Blake Plateau, and Straits of Florida:
*AES Ocean Express. 2002. Application of AES Ocean Express LLC for a natural gas pipeline right-of-way on the outer continental shelf off the coast of Broward County, Florida. Application to MMS. AES Ocean Express.
Agassiz, A. 1869. Preliminary report on the Echini and star-fishes dredged in deep water between Cuba and the Florida Reef, by L. F. de Pourtales, Assist. U.S. Coast Survey. Bull. Mus. Comp. Zool. Harvard 1(9): 253-308.
*Agassiz, L. 1869. Report upon deep-sea dredgings in the Gulf Stream, during the third cruise of the U.S. Steamer BIBB, addressed to Professor Benjamin Pierce, Superintendent U.S. Coast Survey. Bull. Mus. Comp. Zool. Harvard 1(13): 363-386.
*Agassiz, A. 1888. Three cruises of the United States Coast and Geodetic Survey Steamer “Blake”, 1. Bull. Mus. Comp. Zool. Harvard 14: 1-314.
*Anselmetti, F. S., G. Eberli, and Z. Ding. 2000. From the Great Bahama Bank into the Straits of Florida: a margin architecture controlled by sea-level fluctuations and ocean currents. Geological Society of America Bulletin 112: 829-844.
Arendt, M., C. Barans, G. Sedberry, R. Van Dolah, J. Reed, and S. Ross. 2003. Summary of Seafloor Mapping and Benthic Sampling in 200-2000m from North Carolina through Florida, Final Report, Deep-water Habitat Mapping Project, Phase II. South Carolina Dept. of Natural Resources, Charleston, S.C., 156 pp.
*Avent, R.M. and F.G. Stanton. 1975. Submersible reconnaissance and research program. Harbor Branch Foundation, 1975 Annual Report.
*Avent, R.M. and F.G. Stanton. 1979. Observations from research submersible of megafaunal distribution on the continental margin off central eastern Florida, Harnor Branch Foundation Technical Report #25, 40 pp.
*Avent, R.M., F.G. Stanton, and J.K. Reed. 1976. Submersible reconnaissance and research program. Harbor Branch Foundation, Annual Report, 52 pp.
39 *Avent, R., M. King, and R. Gore. 1977. Topographic and faunal studies of shelf-edge prominences off the central eastern Florida coast. Int. Revue ges. Hydrobiol. 62: 185-208.
Ayers, and Pilkey. Piston core and surficial investigations of the Florida-Hatteras slope and inner Blake Plateau. [chapter on corals on Blake Plateau]
*Bailey, Norman and K. Kent. 1982. High-resolution seismic-reflection profiles collected aboard R/V Eastward, cruise ESTW 80-8, over Blake Escarpment. U.S. Geological Survey Open-File Report 82-940. (3 sites east of Florida, 7,170 kn of data; microfilm data available)
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40 *BLM. 1979. South Atlantic OCS Benchmark Program, Outer Continental Shelf Environmental Studies. BLM Contract No. AA550-CT7-2. Texas Instruments Inc. [Cape Fear to Daytona, to 285 m] [Executive summary zeroxed only]
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*Bush, J. 1951. Rock from the Straits of Florida. Bull. Amer. Assoc. Petroleum Geologists 35: 102-107.
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*Cairns, S.D. 1979. The deep-water Scleractinia of the Caribbean Sea and Adjacent Waters. Studies of the Fauna of Curacao and Other Caribbean Islands, No. 180, 341 pp.
*Cairns, S.D. 1986. A revision of the northwest Atlantic Stylasteridae (Coelenterata: Hydrozoa). Smithsonian Cont. to Zoology, No. 418, 131 pp.
*Cairns, S.D. 2000. A revision of the shallow-water azooxanthellate scleractinia of the Western Atlantic. Studies of the Natural History of the Caribbean Region, Vol. 75, 240 pp.
Cairns S.D. and R.E. Chapman. 2002. Biogeographic affinities of the North Atlantic deep-water Scleractinia. In Willison, JHM et al. (eds.), Proc. First Intern. Symp. Deep-Sea Corals. Ecology Action Centre and Nova Scotia Museum, Halifax, pp. 30-57.
41 *Calypso Pipeline. 2001. Geohazards assessment, proposed 24” gas pipeline route, Freeport, Grand Bahama Island to Port Everglades, Florida. Report submitted to Calypso Pipeline LLC, project no. 0401-397. Williamson and Associates, Inc, Seattle, WA and Geoscience Earth and Marine Services, Inc, Houston, TX
Cashman, K.V. and P. Popenoe. 1985. Slumping and shallow faulting related to the presence of salt on the continental slope off North Carolina. Marine and Petroleum Geology 2: 260-?
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Child, C. A. 1998. Nymphon torulum, new species and other Pycnogonida associated with the coral Oculina varicosa on the east coast of Florida, Bulletin of Marine Science 63, 595-604.
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Colquhoun, D.J., Arthur, M.A., Dillon, W.P., Hatcher, R.D., Huddlestun, P.F., Poag, C.W., Valentine, P.C., and Popenoe, P. 1991. Southeastern Atlantic Regional Coast Cross-Section, American Association of Petroleum Geologists, Tulsa.
Continental Shelf Associates. 1979. South Atlantic hard bottom survey. BLM Contract No. AA551-CT8-25, 356 pp. [Charleston to Jacksonville, 44- 194 m depths]
*De Silva, D. 1955. The mystery of the tilefish. Sea Frontiers, May Issue, p. 4-8.
Dillon, W.P. 1981. Regional geology. In Dillon, W. P., ed., Summary report on the regional geology environmental considerations for development, petroleum potential and estimates of undiscovered recoverable oil and gas resources of the United States southern Atlantic continental margin in the area of the proposed Oil and Gas Lease Sale No. 78. U.S. Geological Survey Open-File Report 81-749, p. 6-58e.
Dillon, W.P. (ed.) 1981. Summary report on the regional geology, environmental considerations for development, petroleum potential, and estimates of undiscovered recoverable oil and gas resources of the United States southern Atlantic continental margin in the area of proposed Oil and Gas Lease Sale No. 78. U.S. Geological Survey Open File Report 81-749, 108p.
Dillon, W.P. (ed.). l982. Summary of regional geology, petroleum potential, resource assessment and environmental considerations for oil and gas lease sale area #56: U.S. Geological Survey Open-File Report 82-398, 63 pp.
Dillon, W. P. (ed.). l983. Geology report for proposed oil and gas lease sale no. 90; continental margin off the southeastern United States. U.S. Geological Survey Open-File Report 83-186, 125 p., 2 plates.
42 Dillon, W. P. 1983. Regional geology and petroleum potential. In Dillon, W. P., ed., Geology report for proposed oil and gas lease sale no. 90; continental margin off the southeastern United States. U.S. Geological Survey Open-File Report 83-186, p. 6-84.
Dillon, W. P. 1984. Mineral resources of the Atlantic Exclusive Economic Zone. In Conference Record, Oceans '84, Marine Technology Society and IEEE Ocean Engineering Society, p. 431-437. Reprinted in Champ, M.A., Chmn, 1984, Exclusive Economic Zone Papers, MTS/IEEE, p 72-78.
Dillon, W.P. and Kvenvolden, K.A. ?. Gas hydrates in sea floor sediments off southeastern U.S.: Evidence from seismic reflection and drilling data, Alternative energy source. Methane Hydrates Workshop, Technical Proceedings, Department of Energy, Morgantown, W.Va., DOE- METC 82-49, p. 78-81.
Dillon, W.P. and Max M.D. 2000. Oceanic gas hydrate. In Max, M..D., ed., Natural Gas Hydrate in Oceanic and Polar Environments, p. 61-76, Kluwer Academic Publishers, Dordrecht.
Dillon, W.P. and Max M.D. 2000. The U.S. Atlantic continental margin; the best-known gas hydrate locality, Chapter 13. p. 157-170, In Max, M.D. ed., Natural GasHydrate in Oceanic and Polar Environments, Kluwer Academic Publishers, Dordrecht.
Dillon, W.P. and McGinnis, L.D. 1983. Basement structures indicated by seismic-refraction measurements offshore from South Carolina and adjacent areas. In Gohn, G., ed., U.S. Geological Survey Professional Paper 1313, p. O1-O7.
Dillon, W.P. and Paull, C.K. 1978. Interpretation of multichannel seismic-reflection profiles of the Atlantic continental margin of the coasts of South Carolina and Georgia. U.S.Geological Survey Miscellaneous Field Investigations Map MF-936.
Dillon, W.P. and Paull, C.K. 1980. Summary of development of the continental margin off Georgia based on multichannel and single channel seismic reflection profiling and stratigraphic well data. In Arden, D.D. and Beck, B.F., eds., Symposium on Southeastern Coastal Plain Geology, vol.1, 10 p., 3 fig.
Dillon W.P. and Paull, C.K. 1983. Marine gas hydrates - II: Geophysical evidence. In Cox, J. L. (ed.), Natural Gas Hydrates: Properties, Occurrences and Recovery. Boston, Butterworth Publishers, p. 73-90.
*Dillon, William and P. Popenoe. 1988. The Blake Plateau basin and Carolina trough. Chapter 14. pp. 291- 328, In: R. Sheridan and J. Grow (eds.), The Geology of North America, The Atlantic Continental Margin, U.S. Geological Soc. Am., The Geology of North America, 2 vol.
Dillon, W.P., et al. 1975. Sediments, structural framework, petroleum potential, environmental considerations and operational considerations of the United States South Atlantic outer continental shelf. U.S. Geological Survey Open-File Report 75-411, 262 p., l plate.
43 Dillon, W.P., Sheridan, R.E., and Fail, J.P. 1976. Structure of the Western Blake Bahama Basin as shown by 24 channel CDP profiling. Geology 4: 459-462.
Dillon, W.P., Folger, D.W., Ball, M.M., Powers, R, and Wood, G., Jr. 1978. Summary report of the sediments, structural framework, petroleum potential environmental conditions and operational considerations of the United States South Atlantic continental margin. Prepared for Bureau of Land Management for proposed oil and gas lease sale #54. U.S.Geological Survey Open-File Report 78-594, 39 p.
Dillon, W.P., Klitgord, K.D., and Paull, C.K. 1979. Geologic setting of the COST GE-1 drillsite. In Scholle, P.A., ed., Geological studies of the COST GE-1 well, United States South Atlantic Outer Continental Shelf area. U.S. Geological Survey Circular 800, p. 4-6.
Dillon, W.P., Paull, C.K., Buffler, R.T., and Fail, J.P. 1979. Structure and development of the Southeast Georgia Embayment and northern Blake Plateau: Preliminary analysis. In Watkins, J. S., Montadert, L., and Dickerson, P.W., eds., Geological and Geophysical Investigations of Continental Margins: American Association of Petroleum Geologists Memoir 29, p. 27-41.
Dillon, W.P., Paull, C.K., Dahl, A.G., Patterson, W.C. 1979. Structure of the continental margin near the COST GE-1 well site from a common depth point seismic reflection profile. In Scholle, P.A., ed., Geological studies of the COST GE-1 well, United States South-Atlantic Outer Continental Shelf area: U.S. Geological Survey Circular 800, p.97-107.
*Dillon, W.P., Poag, C.W., Valentine, P.C., and Paull, C.K. 1979. Structure, biostratigraphy and seismic stratigraphy along a CDP seismic profile through 3 drill sites on the continental margin off Jacksonville, Florida. U.S. Geological Survey, Miscellaneous Field Investigation Map MF-1090.
Dillon, W.P., Grow, J.A., and Paull, C.K. 1980. Unconventional gas hydrate seals may trap gas off southeast United States. Oil and Gas Journal, v. 78, no. 1, p. 124, 126, 129-130.
Dillon, W.P., Klitgord, K.D., Paull C.K. and Grow, J.A. 1982. Summary of regional geology. In Dillon, W.P., ed., Summary of regional geology petroleum potential, resource assessment and environmental considerations for oil and gas lease sale area #56: U.S. Geological Survey Open- File Report 82-398, p. 5-20.
Dillon, W.P., Klitgord, K.D., and Paull, C.K. 1983. Mesozoic development and structure of the continental margin off South Carolina. In Gohn, G., ed., U.S. Geological Survey Professional Paper 1313, p. N1-N16.
Dillon, W.P., Popenoe, P., Grow. J.A., Klitgord, K.D., Swift, B.A., Paull, C.K. and Cashman, K.V. 1983. Growth faulting and salt diapirism: Their relationship and control in the Carolina Trough, eastern North America. In Watkins, J.S. and Drake, C.L., eds., Studies in Continental Margin Geology, American Association of Petroleum Geologists Memoir No. 34, p. 21-46.
44 *Dillon, W. P., Paull, C. K., and Gilbert, L. E. 1985. History of the Atlantic continental margin off Florida: The Blake Plateau Basin. In Poag, C. W., ed., Geologic Evolution of the United States Atlantic Margin, Van Nostrand, Reinhold, New York, p. 189-215.
Dillon, W.P., Manheim, F.T., Jansa, L.F., Palmason, G. Tucholke, B.E. and Landrum, R.S. 1986. Resource potential of the western North Atlantic Basin. In Vogt, P.R., and Tucholke, B.E., eds., The Geology of North America, volume M, The Western North Atlantic Regions, Geological Society of America, p 661-676.
Dillon, W. P., Valentine, P. C., and Paull, C. K. 1987. Geology of the Blake Escarpment. NOAA Symposium Series for Undersea Research, vol. 2, no. 2, P 177-190.
Dillon, W.P., P. Valentine, and C. Paull. 1987. The Blake Escarpment- a product of erosional processes in the deep ocean. Symp. Ser. For Undersea Research, NOAA’s Undersea Research Program 2: 177-190.
Dillon, W.P., Schlee, J.S. and Klitgord, K.D. 1988. The development of the continental margin of eastern North America - conjugate continental margin to West Africa. Journal of African Earth Sciences, vol. 7, no. 2, p. 361-367.
Dillon, W.P., Trehu, A.M., Valentine, P.C., and Ball, M.M. 1988. Eroded carbonate platform margin - the Blake Escarpment off southeastern United States. In Bally, A.W., ed, Atlas of Seismic Stratigraphy, American Assoc. Petroleum Geologists Studies in Geology Series, No. 27, vol. 2, p. 40-47.
*Dillon, W.P., Risch, J.S., Scanlon, K.M., Valentine, P.C., and Huggett, Q.J. 1993. Ancient crustal fractures control the location and size of collapsed blocks at the Blake Escarpment, east of Florida. In Schwab, W.C., Lee, H.J. and Twichell, D.C., eds., Submarine Landslides: Selected Studies in the U.S. Exclusive Economic Zone, U.S. Geological Survey Bulletin 2002, p. 54-59.
Dillon, W.P., Fehlhaber, Kristen, Coleman, D.F., Lee, M.W., and Hutchinson, D.R. 1995. Maps showing gas hydrate distribution off the east coast of the United States. U.S. Geological Survey Miscellaneous Field Investigations Map, MF 2268, 2 sheets, 1:1,000,000.
Dillon, W., Hutchinson, D., and Drury, R. 1996. Seismic reflection profiles on the Blake Ridge near Sites 994, 995 and 997. Proceeding of the Ocean Drilling Program, Initial reports, v. 164, p. 47-56
Dillon, W., Holbrook, W.S., Drury, R., Gettrust, J., Hutchinson, D., Booth, J. and Taylor, M. 1997. Faulting of Gas-Hydrate-Bearing Marine Sediments ? Contribution to Permeability. Proceedings of the Offshore Technology Conference, p. 201-209.
45 Dillon, W.P., Danforth, W.W., Hutchinson, D.R., R.M., Drury, Taylor, M.H., Booth, J.S. 1998. Evidence for faulting related to dissociation of gas hydrate and release of methane off the southeastern United States. In Henriet, J.P. and Mienert, J, eds., Gas Hydrates: Relevance to World Margin Stability and Climate Change, Geological Society, London, Spec. Publication 137, p.293-302.
Dillon, W.P., Nealon, J.W., Taylor, M.H., Lee, M.W., Drury, R.M., and Anton, C.H. 2001. Seafloor collapse and methane venting associated with gas hydrate on the Blake Ridge ? causes and implications to seafloor stability and methane release. In C.K. Paull and W.P. Dillon, eds., Natural Gas Hydrates: Occurrence, Distribution, and Detection, American Geophysical Union, Geophysical Monograph 124, p. 211-233.
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Fluke, L.A. 1994. Recent Atlantic shelf sedimentation within a siliciclastic-carbonate transition, Florida, USA. M.S. Thesis, Florida Institute Technology, Melbourne, Florida, 78 p.
46 Folger, D.W., Dillon, W.P., Grow, J.A., Klitgord, K.D., and Schlee, J.S. 1979. Evolution of the Atlantic Continental Margin of the United States. In Talwani M., Hay, W. and Ryan, W.B.F., eds., Deep drilling results in the Atlantic Ocean: Continental Margins and Paleoenvironment. American Geophysical Union, Maurice Ewing Series 3, p 87-108.
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47
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Reed, J.K. 1981e. Research on shelf-edge Oculina coral banks off central eastern Florida. Pp. 32-33, In Summary of Results of Reefs and Hardgrounds Workshop, Duke University Marine Lab, Beaufort, N.C. Sept. 1981.
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Sheridan, R.E., Crosby, J.T., Kent, K.M., Dillon, W.P. and Paull, C.K. l981. The geology of the Blake Plateau and Bahamas region. In: The geologic atlas of the North American borderlands: Canadian Society of Petroleum Geologists, Memoir 7, p. 487-502.
*Siegler, V.B. 1959. Reconnaissance survey of the bathymetry of the Straits of Florida. Univ. Miami, Inst. Mar. Sciences, Marine Lab Rept. 59-3, 9 p.
*Smith, W.H. and D. Sandwell. 1997. Global sea floor topography from satellite altimetry and ship depth soundings. Science 277: 956-1962.
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*Stanley, Daniel. 1969. Atlantic continental shelf and slope of the United States- color of marine sediments. Geological Survey Prof. Paper 529-D, pp. D1-15.
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*Stewart, H.B., Jr. 1962. Oceanographic cruise report, U.S. Coast and Geodetic Survey ship EXPLORER-1960. U.S. Dept. Commerce, Coast and Geodetic Survey, Wash., D.C., 162 p. [parts zeroxed- some stations in Florida Straits]
Sylwester, R.E., Dillon, W.P., Grow, J.A. 1979. Active growth fault on seaward of the Blake Plateau. In Gill, D. and Merriam, D.F., eds., Geomathematical and Petrophysical Studies in Sedimentology: Oxford, Pergamon Press, p. 197-209.
Taylor, M., Dillon, W. Anton, C., and Danforth, W. 1999. Seismic reflection surveys of the Blake Ridge, R/V Cape Hatteras 1992 and 1995: Data acquisition, navigation and processing. U.S. Geological Survey Open File Report 99-372 (2- CD-ROMs).
61 Taylor, M.H., Dillon, W.P., and Pecher, I.A. 2000. Trapping and migration of methane associated with the gas hydrate stability zone at the Blake Ridge Diapir: new insights from seismic data, Marine Geology, vol. 164, p. 79-89.
Teichert, Curt. 1958. Cold- and deep-water coral banks. Bull. Amer. Assoc. Petroleum Geologists 42: 1064-1082.
Tendal, O.S. 1992. The North Atlantic distribution of the octocoral Paragorgia arborea (L., 1758). Sarsia 77: 213-217.
Thompson, M.J. and L.E. Gulliland. 1980. Topographic mapping of shelf edge prominences off southeastern Florida, Southeastern Geology 21, 155-164.
*Top Spot, Inc. 2000? East Florida offshore fishing and diving map, Port Canaveral to Lake Worth Inlet, Map No. N220. Pasadena Hot Spot, Inc., 4016 Strawberry Rd., Pasadena, TX 77504. (bathymetry of Florida Straits)
*Top Spot, Inc. 2000? South Florida offshore fishing and diving map, Miami to Dry Tortugas, Map No. N210. Pasadena Hot Spot, Inc., 4016 Strawberry Rd., Pasadena, TX 77504. (bathymetry of Florida Straits)
Twichell, D.C., Dillon, W.P., Paull, C.K., and Kenyon, N.H. 1996. Morphology of carbonate escarpments as an indicator of erosional processes, Chapt, 6. In Gardner, J.V., Field, M.E., and Twichell, D.C., eds., Geology of the United States' Seafloor: The View from GLORIA, Cambridge University Press, Cambridge, U.K., p. 97-107.
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62 *Uchupi. Elazar and K. Emery. 1967. Structure of continental margin off Atlanic coast of United States. Amer. Assoc. of Petroleum Geologists Bull. 51: 223-234.
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Virden, W. T., T. L. Berggren, T. A. Niichel, and T. L. Holcombe. 1996. Bathymetry of the shelf- edge banks, Florida east coast, National Oceanographic and Atmospheric Administration, National Geophysical Data Center, National Marine Fisheries Service, Beaufort, North Carolina: 1.
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Part II- Gulf of Mexico:
Ballard R. and E. Uchupi. 1970. Morphology and quaternary history of the continental shelf of the Gulf coast of the United States. Bulletin of Marine Science 20: 547-559.
Barry, J., K. Buck, and M. Tamburri. 1998. Cold seep biology and ecology. Monterey Bay Aquarium Research Institute, 1998 Annual Report.
BLM. 1977. Baseline monitoring studies, Mississippi, Alabama, Florida, outer continental shelf, 1975-1976, Vol. 3. Results. Bureau of Land Management Contract No. 08550-CT5-30.
Bright, T.J. ?. Coral reefs, nepheloid layers, gas seeps, and brine flows on hard-banks in the norhteastern Gulf of Mexico. Pp. 40-46, in ?
Bright, T. and L. Pequegnat. 1974. Biota of the West Flower Garden Bank. Gulf Publ. Co., Houston, Tx, 453 p.
Bright, T.J., G. Kraemer, G. Minnery, and S. Viada. 1984. Hermatypes of the Flower Garden Banks, northwestern Gulf of Mexico: a comparison to other western Atlantic reefs. Bull. Mar. Sci. 34: 461-476.
Bright, T., S. Gittings, and R. Zingula. 1991. Occurrence of Atlantic reef corals on the offshore platforms in the northwestern Gulf of Mexico. Northeast Gulf Science 12: 55-60.
Brooks, G. and L. Doyle, 1991. Geologic development and depositional historty of the Florida Middle Ground: a mid-shelf, temperate zone reef system in the northeastern Gulf of Mexico. Society of Sedimentary Geology Special Publication, 46: 189-203.
Brooks and Giammona. 1990. Mississippi-Alabama marine ecosystem study, year 2 annual report. OCS Study, Minerals Management Service 89-0095 to 0096, Contract No. 14-12-0001- 30346, 2 vol.
Brooks, G. and C. Holmes. 1990. Modern configuration of the southwest Florida carbonate slope. Development by shelf margin progradation. Marine Geology 94: 301-315.
Brooks, J., M. Kennicutt, R. Bidigare, and R. Fay. 1985. Hydrates, oil seepage, and chemosynthetic ecosystems of the Gulf of Mexico slope. The Oceanography Report, EOS 66(10): 106.
64 Brooks, J., M. Kennicutt, R. Brigadare, T. Wade, E. Powell, F. Denoux, R. Fay, J. Childress, C. Fisher, I. Rossman, and G. Boland. 1987. Hydrates, oil seepage, and chemosynthetic ecosystems on the Gulf of Mexico slope: an update. The Oceanography Report EOS, May 5, 1987.
Bryant, W., A. Meyerhoff, N. Brown, Jr., M. Furrer, T. Pyle, and J. Antoine. 1969. Escarpments, reef trends, and diapatric structures, eastern Gulf of Mexico. American Association of Petroleum Geologists 53: 2506-2542.
Cairns, S.D. 1977. Stony corals. Mem. of the Hourglass Cruises, Vol. 3, Part 4, 27 pp.
Carsey, J. B. 1950. Geology of Gulf coastal area and continental shelf. Bull. Amer. Assoc. Petroleum Geologists 34: 361-386.
Continental Shelf Associates. 1985. Live bottom survey of drillsite locations in Destin Dome area block 617. Report to Chevron, 40 pp. + photos.
Continental Shelf Associates. 1985. Southwest Florida shelf regional biological communities survey, marine habitat atlas. Minerals Management Service Contract No. 14-12-0001-29036.
Continental Shelf Associates. 1987. Southwest Florida shelf regional biological communities survey, year 3, final report. OCS Study, Minerals Management Service 87-0108 to 0110, Contract No. 14-12-0001-29036, 3 vol.
Continental Shelf Associates. 1992. Mississippi- Alabama shelf pinnacle trend habitat mapping study. OCS Study, Minerals Management Service 92-0026, Contract No. 14-35-0001-30494.
Continental Shelf Associates, Texas A&M University. 1998. Northeastern Gulf of Mexico coastal and marine ecosystem program: ecosystem monitoring, Mississippi/ Alabama shelf, first annual report. OCS Study, Minerals Management Service 97-0037, BRD Contract No. 1445- CT09-96-0006.
Dames and Moore. 1979. The Mississippi, Alabama, Florida OCS baseline environmental survey. MAFLA 1977/1978. Bureau of Land Management Contract No. AA550-CT7-34.
Doyle, L. and C. Holmes. 1985. Shallow structure, stratigraphy and carbonate sedimentary processes of west Florida upper continental slope. American Association of Petroleum Geologists Bulletin 69: 1133-1144.
Environmental Science and Engineering, and LGL Ecological Research Associates. 1985. Southwest Florida shelf benthic communities study. Minerals Management Service Contract No. 14-12-0001-30071, 3 vol.
Environmental Science and Engineering, and LGL Ecological Research Associates. 1986. Southwest Florida shelf benthic communities study, year 5 annual report. OCS Study, Minerals Management Service 86-0074 to 0076, Contract No. 14-12-0001-30211, 3 vol.
65
Environmental Science and Engineering, LGL Ecological Research Associates, and Continental Shelf Associates. 1987. Southwest Florida shelf ecosystems study, data synthesis. Minerals Management Service Contract No. 14-12-0001-30276, 2 Vol.
Florida Institute of Oceonography. 1973. Summary of knowledge of the eastern Gulf of Mexico.
Florida Institute of Oceonography. 1977. Baseline monitoring studies. Mississippi, Alabama, Florida OCS, 1975-1976.
Gittings, S., T. Bright, and E. Powell. 1984. Hard bottom macrofauna of the East Flower Garden brine seep: impact of a long-term sulfurous brine discharge. Contributions Marine Science 27: 105-125.
Gittings, S., T. Bright, W. Schroeder, W. Sager, J. Laswell, and R. Rezak. 1992. Invertebrate assemblages and ecological controls on topographic features in the northeast Gulf of Mexico. Bulletin of Marine Science 50:435-455.
Grassle, J.F. 1985. Hydrothermal vent animals: distribution and biology. Science 229: 713-717.
Hilde, W., G. Sharman, W. Warsi, C. Lee, M. Feeley, and M. Meyer. 1981. Mapping and sub- bottom profiling. p. 6-20, in: Northern Gulf of Mexico Topographic Features Study, Texas A&M Univ.
Holmes, C. 1981. Late neocene and quaternary geology of the southwestern Florida shelf and slope. U.S. Geological Survey, 81-1029.
Hopkins, T., D. Blizzard, S. Brawley, S. Earle, D. Grimm, D. Gilbert, P. Johnson, E. Levingston, C. Lutz, J. Shaw, and B. Shaw. ? A preliminary characterization of the biotic components of composite strip transects on the Florida Middlegrounds, northeastern Gulf of Mexico. U. S. Bureau of Land Management Contract No. 0880-CT5-30, p. 32-37.
Hopkins, T., D. Blizzard, and D. Gilbert. 1977. The molluscan fauna of the Florida Middle Grounds with comments on it’s zoogeographical affinities. Northeast Gulf Science 1(1): 39-47.
Hopkins, T., W. Schroeder, T. Hilde, L. Doyle, and J. Steinmetz. 1981. Northern Gulf of Mexico topographic features study. Bureau of Land Management Contract No. AA551-CT8-35, 150 pp.
Jaap, W. 2000. Observations on deep marine structures: Florida Middle Ground, Pulley Ridge, and Howell Hook from the Deepworker submersible, Sustainable Seas Expedition. Proceedings of American Academy of Underwater Sciences 20th Annual Symposium, St. Petersburg, Florida, October 2000.
66 Jarrett, B., A. Hine, C. Neumann, D. Naar, S. Locker, D. Mallison, and W. Jaap. 2000. Deep biostromes at Pulley Ridge, southwest Florida carbonate platform. Proceedings of American Academy of Underwater Sciences 20th Annual Symposium, St. Petersburg, Florida, October 2000.
Jordan, G. 1951. Continental slope of Apalachicola River, Florida. Bulletin American Association of Petroleum Geologists 35: 1978-1993.
Jordan, G. 1952. Reef formation in the Gulf of Mexico off Appalachicola Bay, Florida. Bulletin Geological Society of America 63: 741-744.
Kennicutt, M.C., J. Brooks, R. Bidigare, R. Fay, T. Wade, and T. MacDonald. 1985. Vent-type taxa in a hydrocarbon seep region on the Louisiana slope. Nature 317: 351-353.
Ludwick, J. 1964. Sediments in northeastern Gulf of Mexico. Pp. 208-238, in: R. Miller (ed.) Papers in Marine Geology, MacMillan Co., N.Y.
Ludwick, J. and W. Walton. 1957. Shelf edge calcareous prominences in the northeastern Gulf of Mexico. Bulletin American Association of Petroleum Petrologists 41: 2054-2101.
Moore, D. and H. Bullis Jr. 1960. A deep water coral reef in the Gulf of Mexico. Bulletin of Marine Science 10: 125-128.
Mullins, H., A. Gardulski, E. Hinchey, and A. Hine. 1988. The modern carbonate ramp-slope of central west Florida. Journal of Sedimentary Petrology 58: 273-290.
Nairn, A. and F. Stehli (eds.). 1975. The Ocean Basins and Margins, V. 3- The Gulf of Mexico and the Caribbean. Plenum Press, N.Y., 706 p.
Neurauter, Thomas. 1980. Bedforms on the west Florida shelf as detected with side scan sonor. M.S. Thesis, Univ. South Florida, 120 p.
Newton C., H. Mullins, F. Gardulski, A. Hine, and G. Dix. 1987. Coral mounds on the west Florida slope: unanswered questions regarding the development of deep-water banks. Palaios 2: 359-367.
Parker, R. and J. Curray. 1956. Fauna and bathymetry of bank on continental shelf, northwest Gulf of Mexico. Bulletin American Association of Petroleum Geologists 40: 2428-2439.
Paull, C.K., B. Hecker, R. Commeau, R. Freeman-Lynde, C. Neumann, W. Corso, S. Golubic, J. Hook, E. Sikes, and J. Curray. 1984. Biological communites at the Florida Escarpment resemble hydrothermal vent taxa. Science 226: 965-967.
Phillips, N., D. Gettleson, and K. Spring. 1990. Benthic biological studies of the southwest Florida shelf. American Zoologist 30: 65-75.
67 Rezak, R., T. Bright, et al. 1981. Northern Gulf of Mexico topographic features study. Bureau of Land Management, Contract No. AA551-CT8-35. [Executive Summary- copied]
Rezak, R. and T. Bright. 1983. Classification and characteristics of banks. Pp. 311-399, in: Reefs and Banks of the Northwestern Gulf of Mexico: their geological, biological, and physical dynamics. Minerals Management Service Contract No. AA851-CT1-55.
Rezak, R., T. Bright, and D. McGrail. 1985. Reefs and banks of the northern Gulf of Mexico: their geological, biological, and physical dynamics. John Wiley and Sons, N.Y., 259 pp. [parts zeroxed] [HBOI Library]
Rezak, R., W. Sager, J. Laswell, and S. Gittings. 1989. Seafloor features on Mississippi- Alabama outer continental shelf. Transactions Gulf Coast Association of Geological Societies 39:511-514.
Rezak, R., S. Gittings, and T. Bright. 1990. Biotic assemblages and ecological controls on reefs and banks of northwest Gulf of Mexico. American Zoologist 30: 23-35.
Rowe, G.T. and D. Menzel. 1971. Quantitative benthic samples from the deep Gulf of Mexico with some comments on the measurement of deep-sea benthos. Bull. Mar. Sci. 21: 556-566.
Schroeder, W., A. Shultz, and J. Dindo. 1988. Inner-shelf hardbottom areas, northeastern Gulf of Mexico. Transactions Gulf Coast Association of Geological Societies 38: 535-541.
Shipp, R. and T. Hopkins. 1978. Physical and biological observations on the northern rim of the Desoto Canyon made from a research submersible. Northeast Gulf Science 2: 113-121.
Uchupi, E. 1967. Bathymetry of the Gulf of Mexico. Gulf Coast Association Geological Society Transactions 17: 161-172.
Uchupi, E. and K. Emery. 1968. Structure of continental margin off Gulf coast of United States. American Association Petroleum Geologists Bulletin 52: 1162-1193.
United States Geological Society. 1998. Geology of shelf-edge habitats of the eastern Gulf of Mexico. USGS Information Handout, Sept. 1998.
Woodward Clyde Consultants and Continental Shelf Associates. 1983. Southwest Florida shelf ecosystems study. Minerals Management Service Contract No. 14-12-0001-29142, 4 vol.
Woodward Clyde Consultants and Continental Shelf Associates. 1985. Southwest Florida shelf ecosystems study. Minerals Management Service Contract No. 14-12-0001-29144, 7 vol.
Part III: Eastern Atlantic and General Deep Sea Reefs:
Fosså, J. H., P. B. Mortensen & D. M. Furevik, 2000a. The deep water coral Lophelia pertusa in Norwegian waters; distribution and fishery impacts. First Internat. Symp. Deep Sea Corals: 25.
68
Fosså, J. H., P. B. Mortensen & D. M. Furevik, 2000b. Lophelia-korallrev langs Nordskekysten forekomst og tilstand. Institute of Marine Research, Bergen, Fisken og Havet Nr. 2: 94 pp.
Fosså, J.H., P.B. Mortensen and D.M. Furevik. 2002. The deep-water coral Lophelia pertusa in Norwegian waters: distribution and fishery impacts. Hydrobiologia 471: 1-12.
Freiwald, A. & J. Schönfeld, 1996. Substrate pitting and boring pattern of Hyrrokkin sarcophaga Cedhagen, 1994 (Foraminifera) in a modern deep-water coral reef mound. Mar. Micropaleon. 28: 199-207.
Freiwald, A., R. Henrich & J. Pätzold, 1997. Anatomy of a deep-water coral reef mound from Stjernsund, west Finnmark, northern Norway. Soc. sedim. Geol., SEPM spec. Pub. 56: 141-161.
Freiwald, A., J. B. Wilson & R. Henrich, 1999. Grounding Pleistocene icebergs shape recent deep- water coral reefs. Sedim. Geol. 125: 1-8.
Genin, A., P.K. Dayton, P.F. Lonsdale, and F.N. Spiess. 1986. Corals on seamount peaks provide evidence of current acceleration over deep-sea topography. Nature 322: 59-61.
Griffin, S. & E. R. Druffel, 1989. Sources of carbon to deep-sea corals. Radiocarbon 31: 533-543.
Hovland, M. and M. Risk. 2003. Do Norwegian deep-water coral reefs rely on seeping fluids? Mar. Geol. 198: 83-96.
Hovland, M., P.B. Mortensen, T. Brattegard, P. Strass and K. Rokoengen. 1998. Ahermatypic coral banks off mid-Norway: evidence for a link with seepage of light hydorcarbons. Palaios 13: 189-200.
Husebo, A., L. Nottestad, J.H. Fossa, D.M. Furevik and S.B. Jorgensen. 2002. Distribution and abundance of fish in deep-sea coral habitats. Hydrobiologia 471: 91-99.
Jensen, A. & R. Frederiksen, 1992. The fauna associated with the bank-forming deepwater coral Lophelia pertusa (scleractinia) on the Faroe shelf. Sarsia 77: 53-69.
Jones, J. B., 1992. Environmenal impact of trawling on the seabed: a review. New Zeal. J. Mar. Freshw. Res. 26: 59-67.
Koslow, J.A. 1997. Seamounts and the ecology of deep-sea fisheries. Amer. Sci. 85: 168-176.
Koslow J.A. and K. Gowlett-Jones. 1998. The Seamount Fauna off Southern Tasmania: Benthic Communities, Their Conservation and Impacts of Trawling. Final Report to Environment Australia and Fisheries Research Development Coorporation. Australia, 104 pp.
69 Koslow, J. A., G. W. Boehlert, J. D. Gordon, R. L. Haedrich, P. Lorance & N. Parin, 2000. Continetal slope and deep-sea fisheries: implications for a fragile ecosystem. ICES J. mar. Sci. 57: 548-557.
Krutschinna, J. & A. Freiwald, 1998. Microendolithic succession along live to dead Lophelia pertusa (L.) skeletons from an aphotic coral reef. Proc. 2nd Inter. Bioerosion Workshop: 43.
Le Goff-Vitry, M.C., O.G. Pybus and A.D. Rogers. 2004. Genetic structure of the deep-sea coral Lophelia pertusa in the northeast Atlantic revealed by microsatellites nad internal transcribed spacer sequences. Molecular Ecol. 13: 537-549.
McDonough, J.J. and K.A. Puglise. 2003. Summary: Deep-sea corals workshop. International planning and collaboration workshop for the Gulf of Mexico and the North Atlantic Ocean. Galway, Ireland, January 16-17, 2003. NOAA Tech. Memo. NMFS-SPO-60, 51 p.
Menzies, R.J., R.Y. George, and G.T. Rowe. 1973. Abyssal Environment and Ecology of the World Oceans. John Wiley and Sons, New York.
Mikkelsen, N., H. Erlenkeuser, J.S. Killingley and W.H. Berger. 1982. Norwegian corals: radiocarbon and stable isotopes in Lophelia pertusa. Boreas 11: 163-171.
Miller, C.A. 2001. Marine protected area framework for deep-sea coral conservation. p 145- 155. In: Willison, J.H.M., J. Hall, S.E. Gass, E.L.R. Kenchington, M. Butler and P. Doherty(eds.). 2001. Proceedings of the First International symposium on Deep-Sea Corals. Ecology Action Centre. Nova Scotia Museum. Halifax, Nova Scotia. 231 p.
Mortensen, P. B. & H. T. Rapp, 1998. Oxygen and carbon isotope ratios related to growth line patterns in skeletons of Lophelia pertusa (L.) (Anthozoa, Scleractinia): implications for determination of linear extension rates. Sarsia 83: 433-446.
Mortensen, P. B., M. Hovland, T. Brattegard & R. Farestveit, 1995. Deep-water bioherms of the scleractinian coral Lophelia pertusa (L.) at 64oN on the Norwegian shelf: structure and associated megafauna. Sarsia 80: 145-158.
Richer de Forges, B., J. A. Koslow & G. C. Poore, 2000. Diversity and endemism of the benthic seamount fauna in the southwest Pacific. Nature 405: 944-947.
Roberts, C.M. 2002. Deep impact: the rising toll of fishing in the deep sea. Trends Ecol. Evol. 17: 242-245.
Roberts, S. and M. Hirshfield. 2003. Deep Sea Corals: out of sight, but no longer out of mind. Oceana. Washington, DC.
Rogers, A.D. 1994. The biology of seamounts. Advances Mar. Biol. 30: 306-350.
70 Rogers, A. D., 1999. The biology of Lophelia pertusa (Linnaeus 1758) and other deep-water reef- forming corals and impacts from human activities. Internat. Rev. Hydrobiol. 84: 315-406.
SGCOR. 2004. Report of the study group on cold-water corals (SGCOR). ICES Advisory Committee on Ecosystems, ICES CM 2004/ACE:07 ref. E.
Squires, D. F., 1964. Fossil coral thickets in Wairarapa, New Zealand. J. Paleontol. 38: 904-915.
Teichert, C., 1958. Cold- and deep-water coral banks. Bull. am. Ass. petrol. Geol. 42: 1064-1082.
Waller R.G. and P.A. Tyler. In Press. The reproductive biology of two deep-sea, reef-building scleractinians from the NE Atlantic Ocean. Coral Reefs.
Wilson, J. B., 1979a. The distribution of the coral Lophelia pertusa (L) [L. prolifera (Pallas)] in the northeast Atlantic. J. mar. biol. Ass. U.K. 59: 149-164.
Wilson, J. B., 1979b. “Patch” development of the deep-water coral Lophelia pertusa (L.) on Rockall Bank. J. mar. biol. Ass. U.K. 59: 165-177.
71 geological with a few biotic observations, mostly observations, biotic few a with geological largely are offU.S. corals southeastern deep the the With the of exception Atlantic. western the in particularly studied, poorly been have habitats coral Deep the StraitsofFlorida. through Carolina South off Plateau Blake the on and Carolina North off occurring sponges) and coral the hard by dominated habitats slope water deeper coral edge off east-central Florida, formed by the stony Two southeastern U.S.: one is located along the shelf depths. these the for reviewed are habitats coral deep general exploiting also now are 2002), development Roberts and exploration 2000; hydrocarbon and al. deeper et into (Koslow rapidly regions expanding are Fisheries 2006). al. et Roberts 2006; al. et Deep (Morgan wide world threats Stream). increasing face ecosystems Gulf coral (i.e., by overlain currents usually extreme are they that fact the and topography bottom rugged their to due partly is recently,until ignored largely this been and have systems reef deep These most waters. U.S. in areas developed, best coral deep 2004) Corcoran and the (Hain extensive have may region This States. United southeastern the off found are habitats coral deep productive and Unique I. 1 2 Washington, DC20013-7012 MRC-153 Smithsonian Inst., P.O. Box37012,NHB, WC-57, Systematics Laboratory Watershed Studies,StPetersburg,FL US GeologicalSurvey, CenterforCoastal & Personnel Act) to: *Currently assigned(throughIntergovernmental Wilmington, NC28409 5600 MarvinMossLn. NationalMarineFisheries ServiceNational UNC-Wilmington,CenterforMarineScience Oculina varicosa Oculina INTRODUCTION Lophelia CAPE HATTERAS TO SOUTHEASTERNFLORIDA Oculina
pu ohr corals other (plus pertusa , STATE OFDEEP CORAL ECOSYSTEMS and the second includes second the and ak, eeecs on references banks, IN THEU.S.SOUTHEASTREGION: STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Steve W. Ross 1 andMarthaS.Nizinski atcnrl lrd. Although off edge Florida. shelf the east-central to distribution in restricted by formed reefs Large Oculina Banks(<150m) Southeastern U.S. History ofDeepCoralResearchoffthe 100 m)areanexception. in this region occur; the Florida bathymetric range where most of the deep corals inhabiting waters deeper than 200 m, which is the depths (deeper m corals emphasize We boundary). EEZ with vary 5000 about to Cape 60 of from range region depth a the and FL, Biscayne, Key to NC, Hatteras, covers chapter and This summarized. corals, briefly are (EEZ) Exclusive Zone U.S. the Economic the in to strategies faunal management associated threats to distributions, assemblages, research coral coral Deep date, U.S. the off southeastern ecosystems coral deep the review We Quattrini 2007). and Ross 2005; al. et Costello 2005; Auster in views conflicting (see opportunistically it occupy to they whether or essential invertebrates or fishes is selected habitat coral deep the whether unclear is it Yet, with habitat. deep-reef unique associated this 2007) closely Quattrini are and invertebrates and (Ross fishes of species off many systems that revealed have coral U.S. southeastern the deep of investigations Our Fish rare. are banks coral deep the to related studies 2004). Mortensen and 1999; Buhl-Mortensen Rogers 1992; Frederiksen harbor and (Jensen reefs deep populations invertebrate species-rich extensive, that revealed Studies elsewhere 2006). al. et Reed 2001; Sedberry in on invertebrates (Reed 2002a, 2002b; references ni te 90 (e rves n ed 2002a, Reed in reviews (see 1960s the until below), these extensive reefs were not described (see corals deep other than m) (60-120 waters were reefs the heavily fished, and are concentrated in shallower structures, substantial forms 2 Oculina Oculina
varicosa . varicosa O. reefs (60- are
233 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
2002b; Reed et al. 2005). The first publication 1980, 1981, 1983), invertebrate communities mentioning these corals along the outer shelf off associated with these reefs (Reed et al. 1982; Florida reported results from seismic transects, Reed and Mikkelsen 1987), Oculina reproduction dredging, and drop cameras (Macintyre and (Brooke and Young 2003, 2005), and fishes Milliman 1970). However, commercial fishermen associated with the Oculina banks (Reed and and Florida scientists apparently knew of these Gilmore 1981; Gilmore and Jones 1992; Koenig reefs earlier (Reed et al. 2005). Macintyre and et al. 2000, 2005; Reed et al. 2006). Milliman (1970) noted Oculina clumps and coral debris along the shelf break from northern Florida These reefs were heavily fished during previous southward and especially from Cape Canaveral to decades and incurred much damage and Palm Beach where ridges in 70-90 m were usually reduction in reef size due to impacts of fishing capped with Oculina. Surveys of these reefs, gear. Research continues, particularly in the using manned submersibles, began in the 1970s zones protected by the South Atlantic Fishery (Avent et al. 1977). Since then, reef monitoring, Management Council (SAFMC). Funding is SOUTHEAST utilizing submersibles and ROVs, has continued lacking, but habitat mapping, restoration, and intermittently (Reed et al. 2005). Additionally, monitoring are high priorities. side-scan sonar surveys were conducted, and multibeam mapping is currently ongoing on these Deep-sea slope corals (>250 m) coral banks (Reed et al. 2005; A.N. Shepard, Historically, deep coral research off the NURC, unpublished data). A variety of studies southeastern U.S. was temporally and spatially have documented coral growth, distributions, sporadic. Until recently deep coral research and upwelling effects on coral growth (Reed was often a by-product of non-coral projects.
Figure 6.1. Southeastern United States regional report area, indicating general areas of Oculina varicosa reefs and the deeper coral (Lophelia mostly) habitats sampled by Ross et al. from 2000-2005 (red stars). The Stetson Bank (white box) is described in the text. Note that these areas do not represent all sites where deep (> 200 m) corals occur nor all sites visited by other researchers. See Reed et al. (2005, 2006) and Partyka et al. (in press) for additional deep coral sites in this region.
234 hy lo eotd pce of species reported also They and corals, hard of species oftwo Banks” major “Stetson occurrence the the confirming 6.1), called (Figure now area this accounting of detailed first the gave (1962) al. et Stetson habitat 1961). (Stetson 1961 in photographed and coral dredged were they until achieved extensive not was these supported that confirmation features However, et (Stetson 1962). mid-1950’s al. the in by sounder surveys depth from resulting Carolina, South off containing topography deep rough corals was discovered on the Blake very Plateau of area An variabilis corals on the Blake Plateau (“Stetson Banks”). In Pratt (1968) presented one photograph of 1968). Uchupi and Zarudzki Tagg1967; Uchupi 1966; and Uchupi (e.g., slope Florida-Hatteras coral the and Plateau termed Blake the on mounds, existed mounds, numerous that noted data echosounding precision on largely based papers geological of series a 1960s the Through al. (1969). et Stetson by described were mounds, coral of hundreds of locations including cruise, 1961 the from details Additional alcyonarians. abundant Caryophyllia profunda Enallopsammia species: Crispatotrochus following the in coral Squires’ collections and re-examined corrected identifications, resulting (1979) Cairns 686 in FL m. Beach, Palm off 1954 collected in dredge species by scleractinian were several noted but (1959) Squires documented. Institution), poorly otherwise (Smithsonian Natural of History Museum National the in deposited Lophelia Some tangles. and trawls beam using 1886 in Plateau Blake the on corals collected Albatross and barren (Agassiz 1888). The research vessel hard being as characterized was Plateau Blake the on bottom the and documented, poorly were steamer theBlake from the of collections 1880 the from resulted Plateau corals of deep report first The roughly (in to beinclusive. reviewed intended not briefly is review this order); are chronological area the in corals deep documented that studies major The Enallopsammia pcmn i toe olcin were collections those in specimens Blake (Agassiz 1888). These collections . and , (=Dendrophyllia (=Caryophyllia Balanophyllia and , Lophelia STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF ohla pertusa Lophelia Bathypsammia s el as well as squiresi ) Tethocyathus )
profunda Lophelia pertusa, . , ,
al. in press), and other mounds may exist. All exist. may mounds other and press), in et al. Partyka 2007; Quattrini and (Ross Carolina North off studied and located been have areas and S.W. Sulak Ross, unpublished data). K.J. Three major coral 1993, Nov (15-18 submersible Navy’s the mounds using survey Coral undersea an during vicinity this found. in located were were coral or bottom hard no but scientist), chief S.W.Ross, cruise, II (R/V 1983 May in sonar by surveyed This published. were data coral McCloskey,no or but Gray,Rowe, Menzies, by directed Lookout Cape a were short other few There comm.). pers. Rowe, G.T. and the and of occurrence “Stetson Banks.” Two of these dives confirmed the DSRV the using dives submersible manned five 1967, of Uchupi 1967) as a result of constantly running constantly of result a as 1967) Uchupi of bank was discovered accidentally (independently coral This cruise. that from is photograph B) 4-4 Figure (1973, al. et Menzies The 1966. June 30 E-4937, 475 m) and dredged (E-4933, 425 m) on (station camera drop by photographed was bank 66, I.E. Gray, chief scientist) during which a coral from a training cruise of the R/V originated mostly records the Carolina North observations, above (1967) Uchupi’s from Aside for (1979) locality Cairns a plotted m. 458 in reef a bottom of a photograph presented and (1969), Menzies and bank off Cape Lookout, repeating “Lophohelia” a figurea inreferenced Rowe vaguely (1973) al. et Menzies data. specific no gave but contour, m Carolinas in “discontinuous banks” along the 450 that suggested Menzies (1968). Rowe and Menzies (1969) later illustrated Roweand by area comment) same without caption the (figure be may which NC, Lookout, Cape off mound coral a of occurrence noted the first (1967) Uchupi profiling, on seismic Based 1960s. late the until off Carolina reported North not were corals such 1962), al. et Stetson 1959; (Squires 1960s early and 1950s late the in investigated and 1880s the in Plateau Blake the on discovered were corals Although 1970), basedon1967 Ball and (Neumann Bay,m) 700-825 FL(around were described from along the slope off Biscayne high) m Additionally,15 (to mounds topped coral s et sudr LR McCloskey (L.R. sounder depth Eastward’s Lophelia tto ae ws rwe and trawled was area station Eastward ee ae n n ra et f the of west area an in made were Alvin in the region (Milliman et al. 1967). al. et (Milliman region the in Enallopsammia cruises to this area off area this to cruises Eastward Lophelia Lophelia Alvin dives. nuclear research nuclear NR-1 p ocre of the off occurred sp. off Cape Lookout. Cape off (=Dendrophyllia Eastward (E-25- Delaware )
235 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
three areas off North Carolina were surveyed (Messing et al. 1990). Genin et al. (1992) noted by multibeam sonar during October 2006 (Ross that sponges and gorgonians were common and Nizinski, unpublished data), revealing many along the outer Blake escarpment (2624-4016 m) mounds that had not been known. The slope off based on observations made during Alvin dives Cape Lookout appears to be the northern extent in 1980. They suggested that these communities of deep corals in the southeast region. were unusually dense for sites lacking sediment. Popenoe (1994) discussed the distribution and Over the next three decades most studies around formation of coral mounds on the Blake Plateau southeastern U.S. deep coral areas continued to and presented a few bottom photographs. Paull be geological and generally not directed toward et al. (2000) surveyed deep-coral habitats off corals. Exceptions include Cairns (1979, 1981, the Florida-Georgia border, dated parts of the 2000, 2001a), who listed ranges for deep sea structures, and suggested that such habitat was Scleractinia and azooxanthellate corals in this very common. Their dating indicated that some region, relying mostly on museum records. mounds may range from 18,000 to 33,000 years SOUTHEAST Neumann et al. (1977) described hard carbonate old. Popenoe and Manheim (2001) extensively mounds in the eastern Straits of Florida off Little reviewed geology, history, and habitats of the Bahama Bank that were covered in various Blake Plateau around the area of the Charleston corals (Lophelia and Enallopsammia) and other Bump, discussing various parameters that may invertebrates. They coined the term “lithoherms” control coral mound formation. Wenner and for these structures. In this same area in 1982, Barans (2001) described benthic habitats of the and also using Alvin, researchers collected and Charleston Bump area and noted some of the aged several “coral” species, indicating that invertebrates and fishes occurring with deep these animals lived from several hundred up to corals. George (2002) discussed a coral habitat, 1800 years (Griffin and Druffel 1989; Druffel et dominated by Bathypsammia tintinnabulum, al. 1990, 1995). These corals have annual rings southeast of Cape Fear, NC (“Agassiz Coral that contain a wealth of information about past Hills”) in 650-750 m. Apparently, the B. climates, ocean productivity, and contamination. tintinnabulum used by Emilini et al. (1978) came This significant discovery has vast implications for from the collections noted by George (2002). the scientific value of deep corals as proxies for A multibeam sonar survey of this site in 2006 climate change and recorders of environmental (Ross and Nizinski, unpublished data) revealed histories (Williams B et al. 2006; Williams et al. a flat bottom with no suggestion of coral mounds. in press). Ayers and Pilkey (1981) documented Reed (2002a, 2002b; Reed et al. 2006) described several coral banks, collected corals, and dated several large areas of deep corals on the Blake some coral samples during a study of sediments Plateau and listed some of the fauna observed. of the Florida-Hatteras slope and inner Blake As part of a SEAMAP bottom mapping project, Plateau. Depending on location in a core, their data and reports to be examined for evidence dead coral samples ranged in age from 5,000 to of deep corals in this area were summarized by 44,000 years old. They dated a living specimen Arendt et al. (2003). This project was completed at 680 years old, but suggested that this age in 2006 and will be incorporated into the South probably reflected the age of the carbon pool in Atlantic Fishery Management Council’s internet the surrounding water. Pinet et al. (1981) also display. mapped coral banks overlapping the same area as Ayers and Pilkey (1981). Blake et al. (1987) Beginning in 2000 and continuing through the briefly mentioned the presence of some soft present, deep coral (or related habitat) research in and hard corals on the Blake Plateau. Many the southeastern U.S. was stimulated by funding deep-reef locations were indicated by the U.S. of studies through the NOAA Office of Ocean Geological Survey sidescan sonar mapping Exploration (see http://oceanexplorer.noaa. (cruises in 1987) of the continental slope (EEZ- gov/explorations) and supplemented by other SCAN 87 Scientific Staff 1991); however, habitats sources. Teams lead by Principal Investigators were not verified in this large scale geological S.D. Brooke, S.A. Pomponi, S.W. Ross, and G.R. survey. Perhaps the first study to document the Sedberry explored deep-coral banks throughout invertebrate community associated with deep- the southeast, mapping habitats, cataloging coral habitat in this region reviewed biozonation fauna, and conducting basic biological studies. of lithoherms in the northeastern Straits of Florida A multi-investigator effort to create detailed
236 ig cnan etnie a hdae deposits hydrate gas extensive contains Ridge and Carolina Rise (Markl et al. 1970). The Blake less steep profile and grades intotypical, the Blakemore Ridge a exhibits Spur Blake the of north Plateau Blake the of slope eastern The Spur). m) on the southeastern margin (south of the Blake the Blake Escarpment (descending to about 4800 Florida-Hatteras Slope (shelf to about 600 m), and major two topographic exhibits breaks, Plateau one on Blake its western The Manheim margin, the and 2001). Popenoe depths, m broad (400-1250 a Stream Gulf Plateau, the by formed Blake feature depositional the by dominated wide and is is unusually It U.S. most configurations. of slope atypical is to Florida) of Carolina the Straits North the of (central much U.S. through southeastern slope continental The rugged topography. a creating and substrates hard exposing often scoured a steep channel along most of its length, bottom topology of the southeast region, and has the shaping force dominant Tertiary)a early the (since been has and is Stream Gulf The 1968). (Pratt topography slump a by characterized and North central From Carolina northward the slope is particularly steep region. southeast the missing in are slope the across cut that canyons major Mexico, of Gulf and Bight Atlantic Middle largely the Unlike a corals). (including fauna of subtropical attachment for al. substrate provide et Avent these and 1980), 1970; Gilliland and Thompson origins 1977; Milliman various and of (Macintyre prominences topographic Florida the of region the including m), (<200 The edge shelf are 1968). (Pratt origin sediments terrigenous of largely whose province sedimentary acarbonate as classified been have slope and and Manheim 2001). The southeastern U.S. shelf Popenoe 1988; Popenoe and Dillon 1979; al. et Schlee 1977; al. et Avent1968; Pratt in reviews (see studied continental well been has rise and U.S. slope shelf, southeastern the of Geology II. by theseefforts. forthcoming from the considerable data collected are publications Future press). in al. et (Partyka U.S. from underway is area the in SEADESC) dives submersible past initiative, (Southeastern Corals Deep-Sea classifications habitat GEOLOGICAL SETTING Oculina ak, s akd y numerous by marked is banks, STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF et al.2005,2006). (Neumann et al. 1977; Messing et corals al. 1990; Reed deep of variety Florida of Straits the of substrates hard Aon occur 2005). al. et (Reed to 400 m 200 that exhibits in varied hardbottom topography platform carbonate a Terrace, Miami the by bordered is Florida of Straits the of side northwestern The 1977). al. et Neumann 1970; Hurley and (Malloy platform carbonate a of part are that lithoherms and mounds ridges, valleys, spaced closely exhibits bottom Stream the and system), Gulf the of (part current Florida the by swept is area This Florida. Key of Straits the offwithin (to region this Biscayne, southern of border for this part report) is southern mostly The mounds (Uchupi1967). Cretaceous to Miocene aged hard substrata, and depressions, numerous exposing Stream, the Gulf by Pleistocene) the during (mostly eroded heavily been has particularly Plateau Blake the of side western The 1972). (Brundage plateau the Manganese over abundant are nodules and pavements 2001). Manheim and (Popenoe scoured by this current, exposing hard substrates flow, Stream heavily is and seaward, Stream Gulf the deflects Gulf to barrier partial a presents the Blake Plateau is the Charleston Bump, which of structure topographic dominant One 1964). Heezen and Pratt 1962; al. et (Stetson mounds the on and ridges and scarps the of edges the in along common are corals occur deep and region, this depressions and plateaus, scarps, Numerous mounds, 1964). Heezen and (Pratt material as well as corals and other invertebrates en cnrbtd y trpd and pteropod by contributed being carbonates with province, carbonate Bahamian the of extension an is Plateau Blake The 2003). al. et Dover van 1997; al. et (Borowski area this in community seep methane known only the and n fclttn hg cret pes u to (up speeds current high northward, facilitating and sediments North southern transporting to Carolina), Florida northern between offshore(mostly to areas delivery sediment offshelf cutting by depths slope continental at even This well studied system influences bottom conditions Current). Florida (or current Stream Gulf the is U.S. outer southeastern the the off slope of and shelf biology and geology the of much The dominant oceanographic feature that shapes III. OCEANOGRAPHIC SETTING Globigerina
237 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
2.5 kn, 125 cm/s) on the bottom (Pratt 1968; particularly lacking. High current speeds have Popenoe and Manheim 2001). The jet-like flow been reported (see above), but currents can vary of the Gulf Stream has a surface width around from near zero to >50 cm sec-1 over short time 80-150 km with a depth of 800-1200 m, fastest scales (pers. obs.). Bottom currents are more current speeds being near the surface center complex around coral mound or rocky features (Bane et al. 2001). The Gulf Stream provides and are accelerated through valleys and over a seasonally stable temperature (generally >27o the tops of mounds/ridges (pers. obs.). Recent C) and salinity (generally >36 psu) regime for the multibeam sonar mapping suggested that long outer shelf and slope. This current transports term current scouring helped shape deep-coral an increasingly massive volume of water as it mounds off North Carolina, but that conditions moves northward (Bane et al. 2001). Inshore of were different at the deeper Stetson area habitats the Gulf Stream (ca. <40 m depth) the physics of (Ross and Nizinski, unpublished data). Bottom the continental shelf water column is dominated temperatures around southeastern U.S. deep by tides, meteorology (e.g., winds, rainfall), and coral habitats (370-780 m) ranged from 5.4° to SOUTHEAST gravity waves (Pietrafesa 1983; Mathews and 12.3° C and salinities varied little from 35 psu Pashuk 1986). (Ross and Quattrini 2007). Similar environmental data from southeastern U.S. deep coral habitats This complex current meanders (influenced in were reported by Reed et al. (2006). part by bottom topography) and produces eddies that spin away from the main current (Atkinson et al. 1985; Bane et al. 2001). As the Gulf Stream IV. STRUCTURE-FORMING DEEP is deflected offshore away from the shelf edge, CORALS OF THE SOUTHEASTERN particularly off South Carolina (by the Charleston U.S. Bump) and off central North Carolina, nitrogen- rich deeper water upwells onto the outer shelf, The southeastern U.S. slope area, including leading to localized areas of enhanced carbon the slope off the Florida Keys, has a unique production (Atkinson et al. 1982; Lee et al. 1991). assemblage of deep-water scleractinians Much of this carbon is subsequently transported (Cairns and Chapman 2001). The warm offshore (Lee et al. 1991). As the Gulf Stream temperate assemblage identified by Cairns and has varied in position since the Pleistocene, it Chapman (2001), encompassing nearly the may alternately uncover deep bottom substrata same geographic range as that covered here, suitable for deep coral settlement or facilitate consists of about 62 species, four of which are burial of coral mounds (Zarudzki and Uchupi endemic to the region. This group of corals was 1968). characterized by many free-living species, few species living deeper than 1000 m, and many Being on the shelf edge, the Florida Oculina species with amphi-Atlantic distributions. For the banks experience more temperate conditions. southeastern U.S., in areas deeper than 200 m, Bottom water temperatures can vary widely over we report a similar assemblage, consisting of 57 short time scales (days to weeks) and on some species of scleractinians (including 47 solitary banks can range from about 7° to 27° C (Avent et and ten colonial structure-forming corals), four al. 1977; Reed 1981), being alternately washed antipatharians, one zoanthid, 44 octocorals, one by warm Gulf Stream or inshore waters and pennatulid, and seven stylasterids (Appendix deeper, cold waters. The colder conditions are 6.1). Thus, the region contains at least 114 usually caused by upwelling, which also provides species of deep corals (Classes Hydrozoa and an increased amount of nutrients (Reed 1983). Anthozoa). We note, however, that this list is Bottom visibility on these banks is generally low, conservative, and we expect that more species current speeds can be variable (sometimes >50 will be discovered in the region as exploration cm sec-1), and sedimentation rates moderately and sampling increase. Since solitary corals do high (15-78 mg cm-2 day-1) (Reed 2002b). not form reefs and are poorly known, we do not treat them in detail. Below we discuss the major While the surface and upper water column structure-forming corals (Appendix 6.1) that most oceanography beyond the shelf edge are fairly contribute to reef-like habitats in the southeastern well studied, bottom conditions over most of the U.S. slope are not well known. Long term data are
238 1986). al. et (Genin recruitment faunal and growth coral column, enhancing the environment for water continued the of physics the alters reef growing the filter-feeding invertebrates and other biota. Thus, substrata, open accelerate from bottom currents which favors m attached 100 mounds, over reef rising and some ridges 1994; The (Rogers 1997). seamounts Koslow to similar ways in productivity local enhance and biota concentrate We hypothesize that high profile deep-coral reefs Figure 6.2. Wilmington. NOAA UnderseaResearchCenteratUNC- the FloridaOculina Selectedphotographs(May2003)from HAPC.Photocredit:L.Horn, STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF oxnhla, u te epr om os not. does form deeper the but zooxanthellae, of form water 70-100 m (Figure 6.1; Reed 2002b). The shallow off east-central Florida, 27 Florida, east-central off reefs large forms only coral this depths, m 2-152 in Caribbean the and Mexico of Gulf the through is south Carolina North m) and Bermuda from occurs it (<200 Although coral). tree shelf (ivory varicosa Oculina outer the U.S. on coral southeastern structure-forming dominant The Scleractinia) a. StonyCorals(Class Anthozoa, Order Oculina a hv symbiotic have may o 32’ N to 28 to 32’N o 59’ N, in 59’N,
239 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
The deeper reefs are almost monotypic mounds the area (especially on the Blake Plateau) seem and ridges which exhibit a vertical profile of 3-35 to form by coral colonization of appropriate hard m (Avent et al. 1977; Reed 2002b). Superficially, substrates, without mound formation by the these structures (Figure 6.2) resemble the deep corals. If bottom currents are too strong, mound reefs formed by Lophelia pertusa. Despite cool formation may be prevented (Popenoe and temperatures, the shelf edge Oculina exhibit rapid Manheim 2001) because sediments cannot be growth, probably facilitated by regular upwellings trapped. Ayers and Pilkey (1981) suggested that of nutrient rich water (Reed 1983). Gulf Stream currents may erode coral mounds, and that present coral bank sizes may be related Lophelia pertusa, the major structure-forming to historical displacements of that current. coral in the deep sea, is the dominant scleractinian Assuming currents also carry appropriate foods, off the southeastern U.S. This species has it may be that currents with variable speeds a cosmopolitan distribution, occurring on the or at least currents of moderate speeds (fast southeastern U.S. slope, in the Gulf of Mexico, enough to facilitate filter feeding but not too fast SOUTHEAST off Nova Scotia, in the northeastern Atlantic, the to prevent sediment entrapment) coupled with a South Atlantic, the Mediterranean, Indian Ocean supply of sediment are the conditions necessary and in parts of the Pacific Ocean over a depth to facilitate coral mound formation (Rogers 1999). range of 50 to 2170 m (Cairns 1979; Rogers Regardless of how coral formations are created, 1999). The 3380 m depth record off New York for we agree with Masson et al. (2003) that elevated L. pertusa reported by Squires (1959) was based topography appears to be an important attribute on a misidentifed specimen (Cairns 1979). Coral for well developed coral communities. habitats dominated by L. pertusa are common throughout the southeastern U.S. from about 370 Deep-coral reefs are fragile and susceptible to to at least 800 m depth. physical destruction (Fossa et al. 2002). It is estimated that these deep reefs may be hundreds Although Lophelia may occur in small scattered to thousands of years old (Neumann et al. 1977; colonies attached to various hard substrata, it Wilson 1979; Ayers and Pilkey 1981; Mikkelsen also forms complex, high profile features. For et al. 1982; Mortensen and Rapp 1998); instance, off North Carolina, Lophelia forms what however, aging data are so limited (especially in may be considered classic mounds that appear the western Atlantic) that age of coral mounds in to be a sediment/coral rubble matrix topped with the western Atlantic is unclear. Recent drilling almost monotypic stands of L. pertusa (Figure on coral mounds off Ireland indicated that these 6.3). Along the sides and around the bases of structures started forming over two million these banks are rubble zones of dead, gray coral years ago and that formation was not related pieces which may extend large distances away to hydrocarbon seeps (Williams T et al. 2006). from the mounds. To the south sediment/coral While the genetic structure (gene flow, population mounds vary in size, and L. pertusa and other relationships, taxonomic relationships) of Lophelia hard and soft corals populate the abundant hard in the northeastern Atlantic is being described substrata of the Blake Plateau in great numbers (Le Goff-Vitry et al. 2004), such studies are just (Figures 6.4 and 6.5). beginning in the western Atlantic (C. Morrison et al. unpublished data). Preliminary genetic Data are lacking on how Lophelia coral banks in results from the southeast region suggest that the the southeastern U.S. are formed. Hypotheses population structure of L. pertusa is more diverse for coral mound formation in the northeastern than expected (C. Morrison et al. unpublished Atlantic were proposed (Hovland et al. 1998; data). Understanding the population genetics Hovland and Risk 2003; Masson et al. 2003), and gene flow will provide insights into coral but it is unclear how relevant these are off the biology, dispersal and distribution of deep corals southeastern U.S. The mounds off North Carolina off the southeastern U.S. and those in other locations off the southeastern U.S. (particularly east of south-central Florida) Although Lophelia is the dominant hard coral off appear to be formed by successive coral growth, North Carolina, other scleractinians contribute collapse, and sediment entrapment (Wilson 1979; to the overall complexity of the habitat (Table Ayers and Pilkey 1981; Paull et al. 2000; Popenoe 6.1). Overall, species diversity of scleractinians and Manheim 2001). Other coral formations in increases south of Cape Fear, NC, but L. pertusa
240 location onthe depthsounderprofiles. Profilesandphotocredit: Rossetal.unpublished data. slender upright growthformof als, anemones, andcongereel(leftphoto), slopecoveredwithanemones andcorals(center photo),andthe illustrates variousliveand dead coralsaswellcoralrubble(centerphoto).Bottom panelshowsmixedcor banks. Top panelshowsliving(white)anddead (gray) coralsnearandonridgetops.Middlepanelalso Figure 6.3 Cape FearLophelia Cape Lookout . Depthsounderprofilesand selectedbottomviewsofthethreeNorthCarolina Lophelia Banks Lophelia Banks A STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Cape Lookout (rightphoto). Habitatphotographsdo not correspondtoaparticular Lopehlia BanksB Lophelia coral -
241 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
Stetson area SOUTHEAST
Jacksonville Banks
South Canaveral
Figure 6.4. Depth sounder profiles and selected bottom views of coral (and sponge) habitats on the Blake Plateau south of North Carolina. Habitat photographs do not correspond to a particular location on the depth sounder profiles. Profiles and photo credit: Ross et al. unpublished data.
242 Madrepora oculata is still dominant. For example, the colonial corals Figure 6.5. lished data. yet beenidentified.Photocredit:Rossetal.unpub south ofNorthCarolina.Othercorals(C)havenot sp.) coralscommonlyfoundontheBlakePlateau toisis spp.,possiblyK.ornata)andgold(E;Gerardia possibly al., unpublished data). et (Ross Florida northern and Carolina South off sites study at habitat reef to adjacent bottom the socialis high. of be aggregations can example, abundance For local region, the of some instances, species particularly in the central portion in But, Most rare. or uncommon either be to appear substrata. hard rubble underlying coral to or attached often U.S. southeastern are the Individuals off found also are 6.1) (Appendix corals solitary the of variety to A adjacent mounds. or on live rather but not to occur singly or as species-specific mounds, south of Cape Fear, NC. These hard corals tend rare off Cape Lookout, NC, are relatively common C A and Leiopathes glaberrima Examplesofblack(A,B;Leiopathes ahpama fallosocialis Bathypsammia and Enallopsammia profunda ), bamboo(D;Kera- STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Thecopsammia Lophelia carpet spp., -
,
r s eaae niis Sm o tee living these of Some colonies entities. separate coral as hard or with association in either singly or in small aggregations, may be observed reach heights of 1-2 m. Black coral colonies, occurring may Colonies region NC. Fear, the Cape to of south limited be to seem distributions their but abundances, moderate in locally occur southeastern and the corals Table6.5; These (Figure 6.1). slope U.S. on corals Leiopathidae structure-forming (Families important are species) four ca. Schizopathidae, corals Black Antipatharia) b. E D B Black corals (Class Anthozoa, Order
243 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
components of the deep reefs attain ages of c. Gold corals (Class Anthozoa, Order hundreds to thousands of years (Williams B et Zoanthidae) al. 2006; Williams et al. in press; C. Holmes and Gerardia spp. colonies are found most often S.W. Ross, unpublished data), and thus, along singly away from other coral structure, but these with gold corals, are among the oldest known corals are also found associated with colonies of animals on Earth. Black corals form annual other structure-forming corals such as Lophelia or regular bands, and these bands contain pertusa, Keratoisis spp., or antipatharians important chemical records on past climates, (Leiopathes spp.). Very little is known about this ocean physics, ocean productivity, pollution, and group of organisms. They apparently exhibit slow data relevant to global geochemical cycles. An growth, reaching ages of at least 1800 years old effort to investigate these geochemical data is (Griffin and Druffel 1989; Druffel et al. 1995) and underway by U.S. Geological Survey (C. Holmes may be valuable in paleoecology studies. and S.W. Ross). SOUTHEAST
Table 6.1. Attributes of structure forming deep-sea corals of the southeastern United States.
Associations with Other Max Structure- Colony Overall Reef- Abun- Colony Morph- Forming Spatial Structural Taxa Building dance Size ology Invertebrates Dispersion Importance Lophelia Yes High Large Branching Many Clumped High pertusa Solenosmilia No Low Small Branching Many Clumped Low variabilis Enallopsammia Low- Small- Low- No Branching Many Clumped profunda Medium Medium Medium Madrepora No Low Small Branching Many Clumped Low oculata Oculina Yes High Large Branching Many Clumped High varicosa Madracis Small- No Low Branching Many Clumped Low myriaster Medium Leiopathes Medium No Medium Branching Many Solitary Medium glaberrima -Large Bathypathes Medium No Low Branching Many Solitary Low alternata -Large Medium No Medium Branching Many Solitary Medium Keratoisis spp. -Large
Table Key Attribute Measure Reef-Building Yes/No Relative Abundance Low/ Medium/ High Size (width or height) Small (< 30cm)/ Medium (30cm-1m)/ Large (>1m) Morphology Branching/ Non-branching Associations None/ Few (1-2)/ Many (>2) Spatial Dispersion Solitary/ Clumped Overall Rating Low/ Medium/ High
244 d. Gorgonians Apni 61. h ms audn species abundant is region the in most observed The 6.1). (Appendix species six only is group this for diversity known the total speciose; is off family No Alcyonacea U.S. southeastern the comprise Nidaliidae, and Nephtheidae, Alcyoniidae, families, Three Alcyonacea) e. True softcorals(Class Anthozoa, Order coral coloniesorasseparateentities. hard with association in either observed be may and aggregations small in or singly either occur colonies coral Bamboo m. 1-2 of heights reach the region south of Cape Fear, NC. Colonies may to limited be to seem also distributions their and 6.1). They occur locally in moderate abundances, distinctive corals and off the southeast region. (Figure 6.5; Table size larger morphology, are also important structure-forming their of because group this of members known best the possibly species), four Isididae, (Family corals Bamboo known onlyfromtheCaribbean). our knowledge of its increased geographic and range (previously species this of specimen fifth known the represented Jacksonville off collected of specimen bipinnata the (Thourella species described newly a offrepresented FL Jacksonville, collected we material date, To region. this for group increase this in species to known of Fear, likely numbers the is Cape sampling Additional of NC. south gorgonians of dramatically diversity increases The 6.1). species 32 (Appendix and represented genera, 17 slope families, seven U.S. by southeastern the on The gorgonians are by far the most diverse taxon Gorgonacea) pce fo fu gnr hv be reported been have Six genera four from region. species southeast the off (Clavulariidae) family one by within represented are (Stolonifera) Alcyonacea, the suborder a Stoloniferans, scleractinian dead as skeletons andcoralrubble. such substrata hard to attaching by habitat the of complexity structural the than diameter, gorgonians. Thus, these and corals add to the overall extent vertical in both size, in smaller are species alcyonacean the of majority The rubble. coral and on dermosponges observed been dead also have to individuals some attached usually is It Florida. off sites at abundant relatively is which (Class Anthozoa, Order hyooga squamata Chrysogorgia Anthomastus agassizi Anthomastus STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF ars 2006); Cairns Lophelia also but , , r kon rm the from known are fishes of species 73 least At Quattrini 2006). in Ross and review (see fauna reef edge shelf shelf edge Oculina the Florida on community fish The Oculina Banks(<150m) V. h Florida The destruction (Koenigetal. 2000). habitat by impacted negatively been also have however,these 1992); Jones and (Gilmore sites aggregation spawning as reefs the use (scamp), groupers, Some microlepis Mycteroperca 2005). 2000, al. et (Koenig destruction habitat caused and complex grouper has the snapper- of reefs members depleted these significantly on fishing commercial years recent In fauna. derived sub-tropically a is this community, invertebrate the like and 2006), al. et Reed 2005; al. et Koenig 1982; SAFMC and increase southwardfromtheCarolinas. stylasterids of diversity and Abundance rubble. often attached to dead scleractinian corals or coral (Appendix 6.1). Individuals observed Individuals 6.1). (Appendix region the from reported been have genera four abundances, representing species great Seven U.S. southeastern in found the off occur commonly corals) (lace stylasterids not Although Anthoathecatae) g. Stylasterids(ClassHydrozoa,Order ( species sertum) isknownintheregion(Appendix6.1). one only museum on records, based and data) unpublished al., et (Ross surveys recent during observed been have pens sea No system. the of diversity and complexity overall the to significantly contributes the southeastern U.S. It is unlikely that this group off pens) (sea pennatulids about known is Little Pennatulacea) f. Pennatulaceans(Class Anthozoa, Order the to southward Caribbean. Carolina North from known are species five other the Atlantic; western the modesta species, Clavularia One 6.1). (Appendix region the from netbae an wt msl sub-tropical ofassociated mostly Densities 6.2). (Figure with affinities fauna invertebrate CORAL COMMUNITIES SPECIES ASSOCIATIONS WITHDEEP banks is typical of the southeastern U.S. Oculina i wdsra throughout widespread is , ef spot diverse a support reefs Oculina gg and (gag) ef (GOMFMC reefs Kophobelemnon M. in situ in phenax are
245 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
invertebrates rival those of shallow coral reef species congregate around deep-coral habitat systems (see review in Reed 2002b). Avent et (Table 6.2). Various crabs, especially galatheoids, al. (1977) presented a preliminary list of benthic are abundant on the deep reefs, playing a role invertebrates dredged from some Oculina of both predator on and food for the fishes. mounds. Analysis of 42 small Oculina colonies Other invertebrates, particularly ophiuroids, yielded about 350 invertebrate species, including populate the coral matrix in high numbers. On 262 mollusc species (Reed and Mikkelson 1987), the relatively barren Blake Plateau, reefs (coral 50 decapod crustacean species (Reed et al. 1982), and hardgrounds) and surrounding coral rubble 47 amphipod species, 21 echinoderm species, 15 habitat seem to offer abundant shelter and food. pycnogoid species, and 23 families of polychaetes (Reed 2002b). The invertebrate community has There are few deep-coral ecosystem references been reduced by habitat destruction (Koenig et al. for the southeast region related to fishes, and 2000). Although Oculina habitats appear to have those are generally qualitative (fishes neither more associated mobile macroinvertebrates than collected nor counted) or fishes were not a specific SOUTHEAST deeper coral areas, large sponges and soft/horny target of the research (Popenoe and Manheim corals are less abundant (Reed et al. 2006). 2001; Weaver and Sedberry 2001; Reed et al. 2005, 2006). In the most detailed study of fishes Deep-sea slope coral areas (>150 m, but most to date, Ross and Quattrini (2007) identified 99 >300 m) benthic or benthopelagic fish species on and Deep coral habitat may be more important to around southeastern U.S. deep-coral banks, 19% western Atlantic slope species than previously of which yielded new distributional data for the known. Some commercially valuable deep-water region. Additional publications resulting from their fish database documented the anglerfish Table 6.2. Dominant benthic fish species (in phylogenetic fauna (Caruso et al. 2007), midwater fish order) observed and/or collected during submersible dives interactions with the reefs (Gartner et al. in (2000-2005) on or near southeastern U.S. Lophelia habitat review), a new species of eel (McCosker based on Ross and Quattrini (2007). Asterisk (*) indicate and Ross in press), and a new species of commercially important species hagfish (Fernholm and Quattrini in press). Common name Although some variability in fish fauna Scientific name (if known) was observed over this region, most of the deep-coral habitat was dominated by Myxinidae (mixed Myxine relatively few fish species (Table 6.2, Figure glutinosa and Eptatretus spp.) hagfishes 6.6). Many of these species are cryptic, Scyliorhinus retifer chain dogfish being well hidden within the corals (e.g., Scyliorhinus meadi Hoplostethus occidentalis, Netenchelys Cirrhigaleus asper roughskin dogfish exoria, Conger oceanicus). Various reef Dysommina rugosa habitats were characterized by Laemonema Synaphobranchus spp. cutthroat eels melanurum, L. barbatulum, Nezumia sclerorhynchus, Beryx decadactylus, Conger oceanicus* conger eel and Helicolenus dactylopterus (Ross Netenchelys exoria and Quattrini 2007). Nearby off reef Nezumia sclerorhynchus areas were dominated by Fenestraja Laemonema barbatulum shortbeard codling plutonia, Laemonema barbatulum, Myxine Laemonema melanurum reef codling glutinosa, and Chlorophthalmus agassizi. Beryx decadactylus usually occurs in Physiculus karrerae large aggregations moving over the reef, Lophiodes beroe while most other major species occur as Hoplostethus occidentalis western roughy single individuals. The morid, Laemonema Beryx decadactylus* red bream melanurum, is one of the larger fishes Helicolenus dactylopterus* blackbelly rosefish abundant at most sites with corals. This fish seems to rarely leave the prime reef area, Idiastion kyphos while its congener L. barbatulum roams Trachyscorpia cristulata Atlantic thornyhead over a broader range of habitats. Although Polyprion americanus* wreckfish Helicolenus dactylopterus (Figure 6.6) can
246 n o te ot mrsie ilgcl aspects biological impressive most the of One (obligate) reeffishes. primary be may habitats deep-coral the around 2007) suggested that some Quattrini of and the fishes (Ross observed Results around habitat. abundant deep-reef is it and substrate, coral the around structures. It is intimately associated often with most occurs it habitats, all in common be Figure 6.6 Photographs credit:S.W. Ross. Helicolenus dactylopterus Laemonema melanurum Polyprion americanus Photographsofsomecommonfishspeciesthesoutheastern USdeep(>200m)coralhabitats (blackbelly rosefish) (reef codling) (wreckfish) STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF hmevs i te ies ad abundant and al. et 2006). Reed diverse and 6.3 (Table fauna the invertebrate is corals the from themselves) (aside habitats coral these of different aspect of the North Carolina deep-coral Carolina North the of aspect different animals or filter food from the currents. One very passing catch to bushes coral on high perched 6.7), (Figure obvious particularly were seastar) and lobster) Hoplostethus occidentalis Eumunida picta Eumunida Beryx decadactylus Conger oceanicus ooii antillensis Novodinia (galatheoid crab; squat crab; (galatheoid (congereel) (westernroughy) (red bream) (brisingid
247 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
habitat compared to the rest of the southeast potential for impacts to deep-sea ecosystems is region is the massive numbers of the brittle star, of great concern because communities at these Ophiacantha bidentata, covering dead coral greater depths are not able to sustain heavy colonies, coral rubble, and to a lesser extent, fishing pressures, as the general longevity of their living Lophelia colonies (Figure 6.7). It is perhaps species, slow growth, and low dispersal rates the most abundant macroinvertebrate on these often prevent recovery from damaging impacts banks and may constitute a major food source for (Koslow et al. 2000; Roberts 2002; Cheung et fishes (Brooks et al. 2007). In places the bottom al. 2007). A large portion of the Oculina banks is covered with huge numbers of several species was closed to fishing due to destruction of habitat of anemones (Figure 6.7). The hydroid fauna is and concern for conservation of corals and the also rich with many species being newly reported associated fauna. There is concern that fisheries to the area and some species being new to may soon target other deep-coral ecosystems in science (Henry et al. in press). The abundance of the region. filter feeders suggests a food rich habitat. Various SOUTHEAST species of sponges, echinoderms, cnidarians Fishing Effects (Messing et al. 1990) and crustaceans (Wenner and Barans 2001) also have been reported from Major human induced damage to habitat and biota deep-coral reefs off Florida, the northeastern has been documented on the east-central Florida Straits of Florida and the Charleston Bump region shelf edge, Oculina reef tract. Extensive damage (Reed et al. 2006). Reed et al. (2006) provided a to corals and fish stocks from fishing operations preliminary list of invertebrates, mostly sponges was reported (Coleman et al. 1999; Koenig et al. and corals, from some deep-coral habitats on the 2000, 2005), including decreased numbers and Blake Plateau and Straits of Florida; however, biomass of corals, decreased amounts of coral most taxa were not identified to species. Lack habitat, and declining fish stocks. The primary fish of data on the invertebrate fauna associated with targets (snapper, grouper, porgy) on the Oculina deep corals is a major deficiency. reefs are also generally considered overfished throughout the waters off the southeastern U.S. Although the invertebrate assemblage associated (SAFMC unpublished data). with northeastern Atlantic Lophelia reefs has been described as being as diverse as shallow On the slope some commercially-exploited water tropical coral reefs (e.g., Jensen and deep-water fishes, like Polyprion americanus Frederickson 1992), data analysis of invertebrates (wreckfish; Vaughan et al. 2001) andHelicolenus associated with western Atlantic deep corals is too dactylopterus (blackbelly rosefish), utilize preliminary to speculate on the degree of species Lophelia habitat extensively (Ross and Quattrini richness. Preliminary data on the invertebrate 2007). Swordfish have been observed along fauna (Nizinski et al. unpublished data) seem the deep reefs (Reed et al. 2006; Ross and to indicate a faunal and habitat transition with Quattrini 2007). Other potentially exploitable latitude. In addition to changes in reef structure species, such as royal red shrimps, rock crabs, and morphology (see above), relative abundance golden crab, squid, bericiform fish species, within a single species decreases, overall species and eels, are also associated with deep-coral diversity increases, and numerical dominance habitats. Signs of past fishing effort (trash, lost between species decreases with decreasing gear) were observed on some banks, but the latitude. In contrast to some fishes, the reef extent to which fishermen sample these areas is associated invertebrate assemblage appears to unknown; therefore, estimations of fishing impact use deep reefs more opportunistically. (Table 6.4) are problematic. The potential for new deep-water fisheries on and around these banks is unknown. At this time our impression is VI. STRESSORS ON DEEP CORAL that benthic fishing impacts to corals and benthic ECOSYSTEMS OF THE fishery species beyond 200 m in this region are SOUTHEASTERN U.S. minimal.
Very little direct information exists to evaluate the health or condition of deep-coral reefs along the coast of the southeastern U.S. However, the
248 are no active offshore production operations. If operations. production offshore active no are there and region, this in exploration hydrocarbon be to appear not issues. do Currently there is these a have federal but moratorium of on habitats, most presently could deep-coral on impacts activities negative anthropogenic Other Effects OfOtherHuman Activities Table 6.3. et al.unpublisheddata,2=Reed2006,3=Henryinreview. southeastern U.S. deep coralhabitats.Coralsarelistedseparatelyin Appendix 6.1. References are1=Nizinski Phylum Arthopoda Phylum Mollusca Phylum Porifera(Sponges) Subphylum Crustacea Class Hexactinellida(glasssponges) Class Malacostraca Class Gastropoda Class Cephalopoda Class Demospongiae Aphrocallistes beatrix multiple species Order Decapoda Coralliophila Squids, Ilexsp. multiple species Octopus, multiplespecies Preliminarylistofdominantbenthicmegainvertebratesobservedorcollectedonnear Infraorder Anomura Other taxa Infraorder Brachyura and theirrelatives) Superfamily Paguroidea(hermitcrabs Family Chirostylidae(squatlobster) Shrimps, multiplespecies Family Portunidae Family Geryonidae Family Pisidae Family Galatheidae(squatlobster) multiple species swimming crab) Munidopsis Uroptychus spp. Gastroptychus salvadori Eumunida picta Bathynectes longispina crab) Chaceon fenneri crab) Rochinia crassa Munida (?)sp. 1,2 1 1
1,2 1,2 spp. including 1 spp. 1 1
Dominant Non-CorallineInvertebrateTaxa 1 STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF
1,2 1,2 1 1 (goldendeepsea 1 (inflated spiny 1 (bathyal 1 Phylum Cnidaria Phylum Echinodermata Phylum Annelida off south Florida could impact deep-coral habitat. pipelines benthic associated with terminal (LNG) U.S. southeastern gas natural liquefied proposed a the of Construction off documented been not has damage such date to to but habitat, damage coral physical cause could laying Cable deep-coral habitat should be carefully considered. to impacts potential the lifted, is moratorium this Class Hydrozoa(Hydroids) Class Echinoidea(seaurchins) Class Anthozoa Class Asteroidea (seastars) Class Crinoidea(crinoids) Class Polychaeta(polychaetes) Class Ophiuroidea(brittlestars) Order Echinoida Order Zoanthidea(zoanthids) Order Actinaria (anemones) Order Cidaroida Order Brisingida(brisingidseastar) Order Echinothurioida multiple species(≥37species) multiple species multiple species multiple speciesincluding multiple species bidentata multiple species rugosa multiple speciesincluding Family Echinidae Family Cidaridae Family Brisingidae Family Echinothuriidae Echinus gracilis Stylocidaris Cidaris rugosa Novodinia antillensis Hygrosoma E. tylodes (Venus flytrapanemone) 1
1
1 1,2 1 spp. spp. , including 1,2 1 1
2 2 1 Eunice
Actinaugi Ophiacantha 3 sp. 1 1
249 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
wall of unidentified anemones Eumunida picta (squat lobster) on Lophelia coral SOUTHEAST
Ophiacantha bidentata (brittle stars) intertwined Novodinia antillensis (brisingid sea star) within the Lophelia coral matrix and Echinus sp. top center.
Bathynectes longispinus Antedonidae (swimming crinoid) (bathyal swimming crab)
Figure 6.7. Photographs of common invertebrates of the southeastern U.S. deep (>200 m) coral habitats. Photo credit: Ross et al. unpublished data.
Bottom disturbance through construction of be in waters shallower than those occupied by offshore tanker ports may impact coral areas, deep corals, designs for deeper water systems especially off Florida where deep water is exist. Coral growth can keep up with a certain closer to shore. Construction of wind farms for amount of sedimentation (Reed 2002b), but high energy production has been recently proposed rates of sedimentation are detrimental to corals for offshore areas. While these would likely (Rogers 1990). We are unaware of references
250 area within a depth range to impact the impact to range depth a within area (Pterois lionfish volitans the species, invasive one only date To impact tocorals(Guinotteetal.2006). oas f h suhat ein (except deep region southeast the to of corals impacts sedimentation documenting Table 6.4.PotentialfishinggearimpactstodeepwatercoralsinthesoutheasternUnitedStates. ae o yt en eotd rm the from reported been yet lionfish not have abundant, seemingly and widespread While 2005). al. et (Meister communities bank atmospheric CO salinity increased and from acidification temperature Ocean conditions. ambient as well as mechanisms, dispersal delivery, food transport, affectsediment could currents these in Changes deep corals. on impacts predict to difficult and Atlantic North conveyor system could overall have far reaching the or current Stream Gulf the of changes that would impact the speed and However,climate direction impact. little have to likely are (increases) level sea in Changes expelled. are unknown, but than those different,to shallow corals where zooxanthellae be azooxanthellate would to corals deep temperature ocean from rising Impacts corals. impacted deep noticeably U.S. southeastern not has change Climate edge orslope. mining of these along the southeastern U.S. shelf current any of unaware are we but manganese), sand, (e.g., area the throughout exist resources this (GOMFMC and SAFMC 1982). Some mineral for potential is there although U.S., southeastern the off activities harvesting coral deep any be to in deep waters of this region. There do not appear absent or rare either be to seem wastes) military Active or municipal, industrial, anthropogenic. (e.g., activities disposal be not usually would Reed 2002b), and if they exist, most such impacts Bottom-set Longline Bottom-set gillnet Mid-water Trawl Traps orPots Bottom Trawl Gear Type Dredge ), has been documented from this 2 is a recently identified potential Severity of Medium Medium Medium Impact High High Low STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Extent of Medium Impact Oculina Oculina Oculina High Low Low Low Low ,
Geographic Extent of UseinRegion VII. a broader appreciation of an ecosystem interactions. incorporates toward which approach management ecosystem-based species single from management snapper/grouper complex. The fishery SAFMC the is of moving part as managed a are wreckfish and plan, through ( regulated crab is deep-sea golden of Harvest waters. slope deeper the in exploited edge shelf the using species to relate dolphin/wahoo and pelagics, coastal complex, snapper/grouper the regulate that Plans plans. management fishery executed through single species or species group (see U.S. fishery southeastern most the of waters and federal in resources habitat of management for responsible is (NMFS), Service Fisheries Marine in cooperation and collaboration with the National The South Atlantic Fishery Management Council, non-fossil corals. in trade international on imposes restrictions CITES Thus, survival. their threaten in specimens of wild animals and plants does not trade international that ensure to is governments between agreement international this of purpose The (CITES). the Flora and Fauna Wild of Species off corals Convention on International Trade in black Endangered and in listed the are of U.S. II southeastern Appendix scleractinian All coral areas. slope deeper the at expected not are they thus, the 2005); al. off et (Meister m 99 depth is U.S. southeastern reported maximum Their area. Low Low Low Low Low Low MANAGEMENT OFFISHERY RESOURCES AND HABITATS Oculina . aaeet is Management www.safmc.net). ak. ee seis are species Fewer banks. Overall Ratingof Gear Impact Medium High Low Low Low Low hco fenneri ) Chaceon
251 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
Swordfish, tunas, sharks and billfishes are or marine sanctuaries. No corals in the area are managed by the Highly Migratory Division of listed as Endangered or Threatened under the NMFS. Endangered Species Act. If other deep-coral reefs prove to be important habitat with a unique Although not applicable to deep corals in this fauna (as they seem to be), these reefs should be region, other species (e.g., sea turtles, whales) considered for protection as are the Oculina coral are protected through such regulations as reefs. There are a variety of potential threats to the Endangered Species Act and the Marine the deep-coral habitats (see above). MPAs or Mammal Protection Act. Sea turtles may occur HAPCs may be viable options for protecting on the Oculina banks; however, most of the slope these systems. However, considerable amounts deep-coral habitat is too deep for sea turtles and and types of data, especially detailed maps, are many marine mammals. In areas to be explored critical for evaluating how and whether to protect for hydrocarbons or mined for minerals within the deep-coral ecosystems (Miller 2001). EEZ, the Minerals Management Service (U.S. SOUTHEAST Dept. of Interior) requires geohazards surveys, The SAFMC is currently evaluating management including documentation of corals, and conducts strategies for southeastern U.S. deep corals. environmental impact reviews of these activities. Considering the needs of the SAFMC to evaluate and manage deep-water habitats in a timely Protection of coral habitat, including deep-water manner, the brief, unpublished descriptions of forms, in this region was established in a Coral, southeastern U.S. deep-coral banks provided by Coral Reef, and Live/Hardbottom Habitat Fishery Ross (2006) and Reed (2004) served as interim Management Plan (FMP) under the Magnuson- tools facilitating potential management options Stevens Fishery Conservation and Management for deep-coral habitats. Based on these reports Act (GOMFMC and SAFMC 1982). This FMP six large areas were recommended as deep coral summarized biological and other data on all HAPCs; these recommendations were modified corals off the southeastern U.S. and in the Gulf in 2006 (Figure 6.8). These proposed HAPC of Mexico. Additionally, optimum harvest of stony areas are included in the current regional FMP corals and sea fans throughout the waters off the and Ecosystem Plan (R. Pugliese, pers. comm.). southeastern U.S. was set at zero (collection for A research plan is being prepared by a SAFMC education and research purposes is permitted). committee to outline gaps in our knowledge and The recent reauthorization of the Magnuson- to address the immediate need for data pertaining Stevens Fishery Conservation and Management to deep-coral habitats on the southeastern U.S. Reauthorization Act (P.L. 109-479) allows continental slope. councils to designate zones for the protection of deep corals and requires research on and monitoring of deep coral habitats. The only deep VIII. REGIONAL PRIORITIES TO coral protected area off the southeastern U.S., UNDERSTAND AND CONSERVE DEEP the Oculina Habitat Area of Particular Concern CORAL COMMUNITIES (HAPC), was described in GOMFMC and SAFMC (1982), but no other deep-coral areas were so Basic data are lacking for the majority of designated. Designation of the Oculina banks as coral habitats >200 m. Recommendations an HAPC became final in 1984, and use of bottom below largely result from basic data needs. disturbing gear was prohibited (Reed 2002b). Considering their habitat value for deep-sea Over the next 14 years, these regulations were communities, their fragility, and a general lack refined and expanded in a series of Amendments of data, locating, describing, and mapping deep to the FMP. Increased protection of the Oculina corals and conducting basic biological studies in banks was granted in 1994, with a total fishing these habitats are global and regional priorities ban within the original HAPC. The HAPC was (McDonough and Puglise 2003; Roberts and doubled in size in 2000, and the new expanded Hirshfield 2003; Puglise et al. 2005). area is now closed to towed bottom gear. In 2004, the ban on fishing was extended indefinitely. Recommendations Detailed mapping of the southeastern U.S. No other deep-coral habitats are designated or shelf edge and slope is critical to better fall within marine protected areas (MPAs), HAPCs understand these habitats and evaluate their
252 the South Atlantic FisheryManagement Council. Figure 6.8. mat ad los rdcin aot the about predictions allows and impacts system facilitates evaluation funds of anthropogenic a in for energy of flow the impact Knowing expended. most column) the provide water would (whole areas and banks surrounding broad coral a of study proposed, trophodynamics be could that studies ecological/biological important many the Of and sites coral areas ofsuspectedcoralmounds. known to given be should the to whole slope off relates the southeastern U.S., priority recommendation this While m. 350-800 of range depth the in possible, especially as soon as conducted be should mapping Multibeam activities. management is the foundation for most other research and contributions to slope ecology. Such mapping Deepcoralareas (redoutlines)proposed forprotectionasHabitat Areas ofParticular Concernby STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF o e gd Acrt got dt o the on data major structure forming growth corals (e.g., Accurate aged. be to Deep corals and the underlying mounds need for needed. done was as fauna, the whole living habitat matrix and associated of documentation Better known. poorly also is fauna deep-coral overall The initiated. be for identification by taxonomic experts, should corals for efforts Collection documentation. better of require region the distributions within corals deep and composition Species Madrepora nomto i ciia t eoytm based ecosystem management. Such to critical is change. information natural of consequences bmo ad lc crl) are corals) black and bamboo , Oculina ef, is reefs, Lophelia ,
253 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
critical to evaluate how banks are formed and and concentrations of reef building deep corals their present status (accreting, eroding). This on the U.S. continental slope. type of research may need to be coupled with local sedimentation and bottom current The three North Carolina Lophelia areas studies. represent the northernmost deep-coral banks off the southeastern U.S. Significant deep- Significant amounts of paleoclimate or coral habitats are not apparent on the U.S. East paleoenvironmental data can be obtained coast again until north of Cape Cod. Because from some coral species. Such studies these banks seem to be a northern terminus for should be pursued. a significant zoogeographic region, they may be unique in biotic resources as well as habitat Genetic studies should continue or be initiated expression. The banks so far examined off North for the major coral species and dominant Carolina are different from much of the coral associated fauna to examine taxonomic habitat to the south on the Blake Plateau. The SOUTHEAST status, dispersal, relationships among coral North Carolina features are dominated by dense banks, and community genetics. thickets of living L. pertusa that cover the tops and sides of the banks; the banks are surrounded If protected areas are established for by extensive coral rubble zones. Unlike areas to southeastern U.S. deep-coral banks, plans the south, the diversity of other corals is low. for long term monitoring, research, education, and enforcement should accompany this Southeastern U.S. deep-coral systems support strategy. The SAFMC is developing such a a well developed community that appears to be plan. Funding should be made available to faunistically different from surrounding non-reef execute the plans. habitats. The fish community on these deep reefs is composed of many species that do not (or at Any deep-water fisheries that currently exist least rarely) occur off the reefs (Ross and Quattrini or that develop on or near the deep-coral 2007). Therefore, they may be considered banks should be carefully monitored and primary reef fishes, in a way similar to those on regulated as deep-water fauna are highly shallow reefs. Many fish species thought to be vulnerable to over fishing, and the habitat is rare and/or outside their reported ranges have subject to permanent destruction. been found on these reefs (Ross and Quattrini 2007). Most likely these species only appeared to be rare because they occurred in areas that IX. CONCLUSION were difficult to sample by conventional means. Thus, these deep-coral habitats support a fish The southeast region contains a huge area of community that appears to be tightly coupled to diverse deep-coral habitat. Rugged topography the habitat and has essentially escaped detection and hard substrata are common on the outer until recently. Invertebrate communities are also shelf edge and slope and this physical structure very diverse and well developed; however, their facilitates development of coral mounds and associations with the reef habitat seem to be other coral habitats. However, detailed maps more opportunistic than is the case for certain fish are lacking, and a major mapping effort must species. However, invertebrate groups are poorly be initiated. Accurate maps are crucial to our known on the slope reefs, and additional data are understanding of the extent of this habitat, for required from diverse habitats to evaluate habitat planning research, and to our ability to manage associations and allow comparisons with other deep-coral habitat. A recent multibeam mapping ecosystems. cruise (Ross and Nizinski unpublished data), covering most of the known North Carolina sites It is clear that the continental slope of the southeast and portions of the Stetson banks, revealed region is important for corals and biodiversity. numerous mounds (probably coral mounds), This is evidenced from the numerous new coral ridges, scarps, and depressions that were habitats discovered, the wide ranging extent and unknown. Based on these and other findings, diversity of corals, numerous species from a it seems probable that the waters off the variety of taxa newly recorded for the area, the southeastern U.S. contains the greatest diversity many species new to science, and the fact that
254 research, isnecessary. ongoing are with coupled protection, Their ecology. slope ecosystems regional of component major these a obviously documented, still is poorly systems of deep-coral biodiversity U.S. while southeastern and 2006), al. et significant (Worm is systems marine in biodiversity of impact overall The value. biomedical significant have may sponges) (e.g., fauna associated or and/ corals Some changes. oceanographic and climate of understanding our increase will that worldwide. Some corals habitats also provide important deep-coral scientific data other any than banks these around recorded were fishes more STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF yr M, iky H 18) itn oe and cores Piston (1981) OH Pilkey MW, Ayers (1977) RH Gore ME, King RM, Avent Auster PJ (2005) Are deep-water corals important (1985) (eds) KA Bush DW,LP, Menzel Atkinson (1982) EE Hofmann LJ, LP,Pietrafesa Atkinson Arendt MD, Barans CA, Sedberry GR, Van Dolah United the of cruises Three (1888) A Agassiz X. ue cnietl hl. SS pn File Open Rept 81-582-A,p5-1-5-89 USGS shelf. continental outer geologic Blake studies on the southeastern Atlantic inner Environmental (ed) P ofthe Popenoe In: Plateau. and slope investigations Florida-Hatteras sediment surficial Hydrobiologie 62:185-208 Gesamten der Revue Internationale coast. Florida eastern central the off prominences Topographic and faunal studies of shelf-edge Springer-Verlag Berlin Heidelberg ecosystems. and in corals 747-760 Cold-water (eds.), Pages JM Roberts A, Freiwald fishes? for habitats Washington, DC156pp Union, Geophysical American Estuarine 2. Sciences and Coastal U.S. Shelf. Southeastern Continental the of Oceanography Marine of Journal Research 40:679-699 Carolina. North Bay, Onslow to sources nutrient of evaluation An Mapping Water Project PhaseII.SAFMC.Charleston,SC Deep Rept Final Florida. through Carolina North from 200-2000m in sampling benthic and mapping seafloor Summary of (2003) SW Ross JK, Reed RF, Zoology atHarvardUniversity14:1-314 Comparative of Museum the of the Bulletin 1. in Mexico, vol 1880. to 1877 from States, United the of of Gulf Caribbean Sea, and along the Atlantic coast the in “Blake” steamer Survey Geodetic and Coast States REFERENCES
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Pratt RM, Heezen BC (1964) Topography of Reed JK, Gilmore RG (1981) Inshore occurrence the Blake Plateau. Deep-sea Research 1 and nuptial behavior of the roughtail 11:721-728 stingray, Dasyatis centroura (Dasyatidae), on the continental shelf, east central Florida. Puglise KA, Brock RJ, McDonough JJ III (2005) Northeast Gulf Science 5:1-4 Identifying critical information needs and developing institutional partnerships to Reed JK, Mikkelsen PM (1987) The molluscan further the understanding of Atlantic deep- community associated with the scleractinian sea coral ecosystems. Pages 1129-1140 in coral Oculina varicosa. Bulletin of Marine Freiwald A, Roberts JM (eds.), Cold-water Science 40:99-131 corals and ecosystems. Springer-Verlag Reed JK, Gore RH, Scotto LE, Wilson KA (1982) Berlin Heidelberg Community composition, structure, aereal Quattrini AM, Ross SW (2006) Fishes associated and trophic relationships of decapods with North Carolina shelf-edge hardbottoms associated with shallow- and deep-water and initial assessment of a proposed marine Oculina varicosa coral reefs. Bulletin of protected area. Bulletin of Marine Science Marine Science 32:761-786 79:137-163 Reed JK, Shepard AN, Koenig CC, Scanlon Reed JK (1980) Distribution and structure of KM, Gilmore RG Jr (2005) Mapping, deep-water Oculina varicosa coral reefs off habitat characterization, and fish surveys central eastern Florida. Bulletin of Marine of the deep-water Oculina coral reef Marine Science 30:667-677 Protected Area: a review of historical and current research. Pages 443-465 in Freiwald A, Roberts JM (eds.), Cold-water corals and ecosystems. Springer-Verlag Berlin Heidelberg
260 ed K Wae D, opn S (2006) SA Pomponi DC, Weaver JK, Reed Ross SW (2006) Review of distribution, habitats, distribution, of Review (2006) SW Ross and reefs coral of Responses (1990) CS Rogers of biology The (1999) AD Rogers seamounts. of biology The (1994) AD Rogers Roberts S, Hirshfield M (2003) Deep Sea Corals: Roberts CM (2002) Deep impact: the rising toll of Roberts JM, Wheeler AJ, Freiwald A (2006) Reefs Rowe GT, Menzies RJ (1969) Zonation of large of Zonation (1969) RJ GT,Menzies Rowe bottom Deep (1968) RJ Menzies GT, Rowe fauna fish The (2007) AM Quattrini SW, Ross tatc ihr Mngmn Council, Management Charleston, SC.2 Fishery Atlantic South to Rept Unpublished FL). Canaveral, Cape to Carolina States (North United slope continental southeastern the on reefs coral water deep of fauna associated and Marine sedimentation. Ecology ProgressSeries62:185-202 to organisms reef of Hydrobiology 84:315-406 Review International activities. human from impacts and corals reef-forming deep- water other and 1758) (Linnaeus pertusa Advances inMarineBiology30:306-350 mind. of out longer no Oceana, Washington, DC,USA ,16pp but sight, of out And Evolution17:242-245 Ecology In Trends sea. deep the in fishing 547 312:543- Science ecosystems. coral water of the Deep: the biology and geology of cold- 375 Science78:343- Marine of Bulletin Mexico. Blake Plateau,Straits of Florida, andGulf of pertusa deep-water of fauna and Habitat Carolinas. Deep-SeaResearch 16:531-537 benthic invertebrates in the deep-sea off the Carolina. North of Deep-Sea Research15:711-719 coast the off currents Deep-Sea States. Research I54:975-1007 United the off southeastern banks coral deep with associated coral reefs off the southeastern U.S.: nd Ed,37pp STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Lophelia Lophelia a Dvr L hrn , enad M Caylor JM, Bernhard P, Aharon CL, Dover Van the of Microrelief (1966) AR Tagg E, Uchupi south margin continental The (1967) E Uchupi Topographic (1980) LE Gilliland MJ, Thompson (1969) JD Milliman E, Uchupi TR, Stetson Coral (1962) RM DF,Pratt Squires TR, Stetson Stetson TR (1961) Report on Atlantis cruise # 266, collected corals sea Deep (1959) DF Squires Stream: the in Island (2001) (ed) GR Sedberry WP,Structure JADillon (1979) Grow JS, Schlee Sea Research I50:281-300 chemosynthetically based ecosystem. Deep- soft-sediment, a of characterization seeps: methane Ridge Blake (2003) R Vrijenhoek Shank BK, Gupta Sen RD, T, TurnipseedM, Seitz JL, Salerno C, Ruppel EC, Raulfs S, W, Gilhooly Goffredi SK, Knick W,KE, Macko SA, Rapoport Flickinger M, Doerries E, of Society Geological America Bulletin77:427-430 Carolina. North Lookout, Cape of south margin continental structure. SoutheastGeology8:155-177 shallow Carolina: North Hatteras, Cape of 21:155-164 off prominences edge southeastern Florida. Southeastern shelf Geologyl of mapping Coast Assoc GeolSoc19:131-142 small areas of the Blake Plateau. Trans Gulf two of morphology subsurface and Surface Novitates Museum 2114:1-39 American Plateau. Blake the on water deep in occurring banks Institute RefNo61-35 June-July 1961. Woods Hole Oceanographic No 1965:1-42 I. Atlantic corals. American Museum Novitates Observatory. Geological Lamont the by the Fisheries pp of American 240 MD, Bethesda, 25, Symposium Society fisheries Bump. Charleston and oceanography Publication 27, Tulsa, OK,pp95-117 Special Mineralogists and Paleontologists Economic of Society Slopes. Continental of States. In: Doyle, LJ, Pilkey OH (ed) Geology of the continental slope off the eastern United
261 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
Vaughan DS, Manooch CS III, Potts JC (2001) HK, Micheli F, Palumbi SR, Sala E, Selkoe Assessment of the wreckfish fishery on the KA, Stachowicz JJ, Watson R (2006) Impacts Blake Plateau. pp 105-119 In: Sedberry GR of biodiversity loss on ocean ecosystem (ed) Island in the Stream: oceanography and services. Science 314:787-790 fisheries of the Charleston Bump. American Fisheries Society Symposium 25, Bethesda, Zarudzki EFK, Uchupi E (1968) Organic reef MD alignments on the continental margin south of Cape Hatteras. Geological Society of Weaver DC, Sedberry GR (2001) Trophic America Bulletin 79:1867-1870 subsidies at the Charleston Bump: food web structure of reef fishes on the continental slope of the southeastern United States. pp 137-152 In: Sedberry GR (ed) Island in the Stream: oceanography and fisheries of
SOUTHEAST the Charleston Bump. American Fisheries Society Symposium 25, Bethesda, MD
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Williams B, Risk MJ, Ross SW, Sulak KJ (2006) Deep-water Antipatharians: proxies of environmental change. Geology 34:773-776
Williams B, Risk MJ, Ross SW, Sulak KJ (in press) Stable isotope records from deep- water Antipatharians: 400-year records from the south-eastern coast of the United States of America. Bulletin of Marine Science
Williams T, Kano A, Ferdelman T, Henriet J-P, Abe K, Andres MS, Bjerager M, Browning EL, Cragg BA, De Mol B, Dorschel B, Foubert A, Frannk TD, Fuwa Y, Gaillot P, Gharib JJ, Gregg JM, Huvenne VAI, Leonide P, Li X, Mangelsdorf K, Tanaka A, Monteys X, Novosel I, Sakai S, Samarkin VA, Sasaki K, Spivack AJ, Takashima C, Titshack J. (2006) Cold-water coral mounds revealed. EOS 87:525-526
Wilson JB (1979) “Patch” development of the deep-water coral Lophelia pertusa (L.) on Rockall Bank. Journal of the Marine Biological Association of the United Kingdom 59:165-177
Worm B, Barbier EB, Beaumont N, Duffy JE, Folke C, Halpern BS, Jackson JBC, Lotze
262 Appendix 6.1. Checklist of deep corals occurring off the southeastern United States (Cape Hatteras, NC to Key Biscayne, FL) at 200-1000 m depth (except shallower Oculina). Higher taxa are in phylogenetic order; families, genera and species are in alphabetical order. Some species have cosmopolitan distributions; however, only the northwestern Atlantic portion of their geographic ranges are reported. MR = museum records (holdings of the National Museum of Natural History, Smithsonian Institution). C & B = Cairns & Bayer (2002, 2003, 2004a, 2004b) *** = Cairns (1979, 2000), Cairns et al. 1999, Cairns & Chapman 2001, unpublished records. S = azooxanthellate solitary scleractinian corals. State of knowledge for the solitary corals is limited; therefore, species- specific geographic and bathymetric ranges are not given. Species included in this list have either been reported from the southeastern U.S. or are likely to occur in the region based on Cairns (1979, 2000), Cairns et al. 1999, and data obtained from unpublished records.
Higher Taxon Species Distribution Depth Range(m) Reference Phylum Cnidaria Class Anthozoa
Subclass Hexacorallia REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Order Scleractinia Family Anthemiphylliidae Anthemiphyllia patera Pourtalès, 1878 S Family Caryophylliidae Anomocora fecunda (Pourtalès, 1871) S Asterosmilia marchadi (Chevalier, 1966) S Asterosmilia prolifera (Pourtalès, 1871) S
Caryophyllia ambrosia caribbeana Cairns, 1979 S Caryophyllia antillarum Pourtalès, 1874 S Caryophyllia berteriana Duchassing, 1850 S Caryophyllia polygona Pourtalès, 1878 S Cladocora debilis Milne Edwards & Haime, 1849 S Concentrotheca laevigata (Pourtalès, 1871) S Crispatotrochus squiresi (Cairns, 1979) S Dasmosmilia lymani (Pourtalès, 1871) S Deltocyathus agassizii Pourtalès, 1867 S Deltocyathus calcar Pourtalès, 1874 S Deltocyathus eccentricus Cairns, 1979 S Deltocyathus italicus (Michilotti, 1838) S Deltocyathus moseleyi Cairns, 1979 S Deltocyathus pourtalesi Cairns, 1979 S
263 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION S S S S S S S S S S S S S S S S S S *** *** *** *** *** *** Reference
800 55-366 146-505 95-2000; 403-1748 220-1383 300-1646 Depth Range commonly 500- SOUTHEAST Distribution Straits of FL; northern Gulf Arrowsmith Bank, of Mexico; Yucatan - Straits of Florida MA Nova Scotia - FL Straits; eastern Nova Scotia - FL Antilles Gulf of Mexico; Lesser Arrowsmith Bank, NC, GA; Yucatan
GA - Suriname GA GA; off Nicaragua GA; off Cairns, 1977 (Pourtalès, 1868) (Chevalier, 1966) (Chevalier, (Pourtalès, 1867) Squires, 1959 (Pourtalès, 1868) (Moseley, 1876) (Moseley, Cairns, 1977 (Pourtalès, 1868) Cairns, 1979 Cairns, 1979 (Duncan, 1873) (Pourtalès, 1867) (Pourtalès, 1871) (Esper, 1794) (Esper, (Pourtalès, 1878) (Pourtalès, 1878) Pourtalès, 1874 (Philippi, 1842) Cairns, 1979 Pourtalès, 1868 Duncan, 1873 (Milne Edwards & Haime, Species (Linnaeus, 1758) Desmophyllum dianthus Labyrinthocyathus facetus Labyrinthocyathus langae Lophelia pertusa Oxysmilia rotundifolia Paracyathus pulchellus Premocyathus cornuformis Solenosmilia variabilis Stephanocyathus coronatus Stephanocyathus diadema Stephanocyathus laevifundus Stephanocyathus paliferus cylindraceus Tethocyathus recurvatus Tethocyathus variabilis Tethocyathus rawsonii Trochocyathus Balanophyllia floridana Bathypsammia fallosocialis Bathypsammia tintinnabulum Cladopsammia manuelensis Eguchipsammia gaditana Enallopsammia profunda Enallopsammia rostrata 1848) Balanophyllia cyathoides Higher Taxon
Family Dendrophylliidae
264 Higher Taxon Species Distribution Depth Range Reference Thecopsammia socialis Pourtalès, 1868 S Family Flabellidae Flabellum atlanticum Cairns, 1979 S Flabellum moseleyi Pourtalès, 1880 S
Javania cailleti (Duchassaing & Michelotti, 1864) S Polymyces fragilis (Pourtalès, 1868) S Family Fungiacyathidae Fungiacyathus symmetricus (Pourtalès, 1871) S Family Guyniidae Pourtalocyathus hispidus (Pourtalès, 1878) S STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Schizocyathus fissilis Pourtalès, 1874 S Stenocyathus vermiformis (Pourtalès, 1868) S NC - FL; Greater Antilles; western 53-801; commonly Family Oculinidae Madrepora carolina (Pourtalès, 1871) Caribbean; Gulf of Mexico 200-300 *** GA - Rio de Janeiro, Brazil; Gulf Madrepora oculata Linnaeus, 1758 of Mexico 144-1391 *** Oculina varicosa Lesueur, 1821 NC - FL; West Indies; Bermuda 3-150 *** Madracis myriaster (Milne Edwards & Haime, GA - Suriname; throughout the Family Pocilloporidae 1849) Caribbean and Gulf of Mexico 20-1220 *** Family Turbinoliidae Cryptotrochus carolinensis Cairns, 1988 S Deltocyathoides stimpsonii (Pourtalès, 1871) S Order Antipatharia GA; FL; Gulf of Mexico (FL;AL; 20 MR; LA); Jamaica; Campeche Bank, Ross et al. Family Leiopathidae Leiopathes glaberrima (Esper, 1788) Mexico; Venezuela 37; 220-685 unpub. Leiopathes spp. 3 MR; SC; GA; FL; Yucatan Channel Ross et al. Family Schizopathidae Bathypathes alternata Brook, 1889 (off Arrowsmith Bank) 412-658 unpub. Parantipathes sp. Order Zoanthidae Family Gerardiidae Gerardia spp.
265 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION
MR 9 MR 1 MR 42 MR 63 MR 44 MR 27 MR 41 MR 25 MR 19 MR unpub. unpub. unpub. Cairns, 28 MR; 47 MR; 218 MR 2001b; 14 Ross et al. Ross et al. Ross et al. Reference
805 1153 2919 1023 29-861 13-105 24-134 24-329 175-586 320-3186 320-1354 430-1050 2107-3506 91-368; 770; 60-878; 1153- Depth Range 137-457; 750- 27-298; 1023- 14-159; 350-400 SOUTHEAST Distribution Canada (off Nova Scotia, Canada (off FL Newfoundland); MA; DE; VA; FL; Gulf of Mexico (FL; TX) FL; Gulf of Mexico (FL; Canada (Nova Scotia, Newfoundland); MA; DE; GA; FL; Bahamas SC; Bahamas; Dominican Republic; Martinique NC; SC; GA; FL FL, SC; FL; Gulf of Mexico (off LA) FL GA; FL; Bahamas; Straits of FL Gulf of Mexico Key West); (off Antilles; Brazil Keys); Lesser (FL Jacksonville, Fl; Caribbean Canada (off Nova Scotia, Canada (off Newfoundland); ME; MA; SC; GA; FL
Canada (off Nova Scotia, Canada (off NC Newfoundland); MA; VA; FL; Bahamas; Colombia; Tobago; Trinidad; Venezuela; Suriname; Dominican Republic; St. Lucia Havana, (off NC; Straits of FL Cuba); Bahamas SC; GA; FL; Bahamas; Straits of Keys; Havana, Cuba); FL (off FL Keys) FL Gulf of Mexico (off Verrill, 1878 Verrill, (Verrill,1883) (Deichmann, 1936) Deichmann, 1936 Verrill, 1922 Verrill, (Pourtalès, 1867) Species (Deichmann, 1936) (Verrill, 1874) (Verrill, (Sars, 1890) Deichmann, 1936 Dana, 1846 (Pourtalès, 1868) Verrill, 1922 Verrill, Bayer, 1961 Bayer, Gersemia fruticosa Siphonogorgia agassizii Anthomastus agassizi Anthomastus grandiflorus Bellonella rubistella Scleranthelia rugosa fruticulosa Telesto nelleae Telesto sanguinea Telesto rudis Trachythela Pseudodrifa nigra Chrysogorgia multiflora Chrysogorgia squamata Clavularia modesta Higher Taxon Family Nephtheidae Order Gorgonacea Family Nidaliidae Order Alcyonacea Order Alcyoniidae Family Family Chrysogorgiidae Family Clavulariidae Subclass Octocorallia
266 Higher Taxon Species Distribution Depth Range Reference Family Coralliidae Corallium sp. Corallium niobe Bayer, 1964 off Jupiter Inlet; Bahamas 659-677; 1023 3 MR Reed 2004; Savannah lithoherms, GA; east Ross et al. Family Gorgoniidae Eunicella modesta (Verrill, 1883) coast FL Lophelia reefs 518-732 unpub. Hudson Canyon; SC; FL; Bahamas; Gulf of Mexico (FL; Family Isididae Acanella eburnea (Pourtalès, 1868) LA; TX); Caribbean Sea (Nevis) 309-2100 34 MR GA; FL; eastern Gulf of Mexico; STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Bahamas; Campeche Bank, Mexico; Guadeloupe; Colombia; Keratoisis flexibilis (Pourtalès, 1868) Venezuela 170-878 29 MR Canada (off Nova Scotia, 45 MR; Newfoundland); MA; GA; FL; Ross et al. Keratoisis ornata Verrill, 1878 Bahamas; Cuba 274-3236 unpub. Lepidisis longiflora Verrill, 1883 FL; Caribbean Sea (Nevis) 743-1125 2 MR Family Paragorgiidae Paragorgia arborea (Linnaeus, 1782) Canada; MA; NJ; MD; VA; NC 247-680 9 MR Florida Straits (off Palm Beach); Paragorgia johnsoni Gray, 1862 Bahamas 522-608 6 MR Canada (off Nova Scotia); GA; FL; Straits of FL (off Havana, Family Plexauridae Paramuricea placomus (Linnaeus, 1758) Cuba) 247-805 6 MR Paramuricea sp. MA; SC; GA; FL; Straits of FL (off FL Keys; Havana, Cuba); Bahamas; Gulf of Mexico (FL; LA); Yucatan Channel (off Swiftia casta (Verrill, 1883) Arrowsmith Bank) 40-1953 49 MR GA; FL; Straits of FL (off FL Keys); Bahamas; Puerto Rico; Gulf of Mexico (FL; MS); Mexico; Panama; Colombia; Venezuela; Tobago; French Guiana; Guyana; Swiftia exserta (Ellis & Solander, 1786) Brazil 18-494 54 MR
267 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION 3 MR 2004a 2004b 2004b 2004a 2004b 2004b Bayer, Bayer, C & B, C & B, C & B, C & B, C & B, C & B, Reference 2001; 8 MR Bayer, 2001 Bayer, C & B, 2002 C & B, 2003 C & B, 2002 C & B, 2003 C & B, 2003 82-514 593-911 183-732 229-556 221-858 161-792 374-555 310-878 348-572 165-706 677-900 738-1473 494-1065 514-2063 Depth Range SOUTHEAST Distribution SC; GA; Bahamas Straits of Florida; Lesser Antilles Straits of Florida; Lesser Straits of FL Straits of FL (off Cape Canaveral (off Straits of FL - Cuba); Bahamas; Lesser Antilles Lydonia Canyon; FL; Gulf of Lydonia Keys) FL Mexico (off central Florida; Bahamas; off Honduras; northern Antilles; off Gulf of Mexico Delray Beach); (off Straits of FL Antilles Bahamas; Lesser Delray Beach); (off Straits of FL Bahamas; Cuba; Puerto Rico; Campeche Bank, Mexico SC to Cuba off North and South Carolina off Insular side Straits of FL (off (off Insular side Straits of FL Palm Beach, north of Little Bahama Bank), Bahamas to Channel Yucatan SC to FL off New England seamounts; Bermuda; eastern coast FL; Antilles; northern Gulf Bahamas; of Mexico St. (off Bermuda; Straits of FL Lucie Inlet, Palm Beach, Delray Beach;Bahamas); Cuba Cairns & Cairns & Bayer, 2004 Cairns & Bayer, Cairns & Bayer, 2004 Cairns & Bayer, Cairns & Bayer, 2004 Cairns & Bayer, Bayer, 2001 Bayer, Cairns & Bayer, 2004 Cairns & Bayer, (Pourtalès, 1868) Species (Johnson, 1862) (Milne Edwards & Haime, Deichmann, 1936 (Kukenthal, 1915) (Deichmann, 1936) (Hickson, 1909) (Wright & Studer, 1889) & Studer, (Wright Calyptrophora trilepis Callogorgia americana 2002 Bayer, Calyptrophora gerdae Swiftia koreni Callogorgia gracilis 1857) Candidella imbricata Narella bellissima Narella pauciflora Narella versluysi Paracalyptrophora duplex Paracalyptrophora simplex Plumarella aurea Plumarella dichotoma Plumarella laxiramosa Higher Taxon Family Primnoidae
268 Higher Taxon Species Distribution Depth Range Reference off NC, through Straits of FL; C & B, Plumarella pellucida Cairns & Bayer, 2004 Bahamas 549-1160 2004b off NC, through Straits of FL; C & B, Plumarella pourtalesii (Verrill, 1883) Cuba; Bahamas 183-882 2004b Cairns, 2006; off northern FL; Straits of FL; off Ross et al. Thouarella bipinnata Cairns, 2006 Little Bahama Bank; off Guyana 507-1000 unpub. Order Pennatulacea STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Family Kophobelemnidae Kophobelemnon sertum Verrill, 1885 off NC 1542 1 MR Class Hydrozoa Order Anthoathecatae Suborder Filifera Cairns, Family Stylasteridae Crypthelia floridana Cairns,1986 eastern, southwestern FL 593-823 1986 GA; Straits of FL (off FL Keys); SE Gulf of Mexico; Yucatan Cairns, Distichopora foliacea Pourtalès, 1868 Channel (off Arrowsmith Bank) 183-527 1986 73-922; commonly Cairns, Pliobothrus symmetricus Pourtalès, 1868 SC through Lesser Antilles 150-400 1986 GA; Bahamas; Yucatan Cairns, Stylaster complanatus Pourtalès, 1867 Peninsula; Virgin Islands 183-707 1986 SC - SW FL; Bahamas; Cay 146-965; Sal Bank; Yucatan Channel (off commonly 650- Cairns, Stylaster erubescens Pourtalès, 1868 Arrowsmith Bank) 850 1986 123-759; SC; Bahamas; Cuba; Yucatan commonly 300- Cairns, Stylaster laevigatus Cairns, 1986 Channel (off Arrowsmith Bank) 400 1986 SC; Straits of FL (off FL Keys); Cairns, Stylaster miniatus (Pourtalès, 1868) Bahamas; Cuba 146-530 1986
269 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION SOUTHEAST
270 ARTICLE IN PRESS
Deep-Sea Research I 54 (2007) 975–1007 www.elsevier.com/locate/dsri
The fish fauna associated with deep coral banks off the southeastern United States
Steve W. RossÃ,1, Andrea M. Quattrini
University of North Carolina Wilmington, Center for Marine Science, 5600 Marvin K. Moss Lane, Wilmington, NC 28409, USA
Received 2 December 2006; received in revised form 12 March 2007; accepted 14 March 2007 Available online 11 April 2007
Abstract
Deep-sea or cold-water corals form substantial habitat along many continental slopes, including the southeastern United States (SEUS). Despite increasing research on deep coral systems and growing appreciation of their importance to fishes, quantitative data on fish communities occupying these ecosystems are relatively lacking. Our overall goals were to document the fish species and their relative abundances and to describe the degree of general habitat specificity of the fishes on and around deep coral habitats on the SEUS slope. From 2000 to 2006, we used the Johnson-Sea-Link (JSL) submersible (65 dives, 366–783 m), supplemented with otter trawls (33 tows, 365–910 m) to document fishes and habitats from off North Carolina to east-central Florida. Eight areas with high concentrations of deep-sea corals were surveyed repeatedly. Three general habitat types (prime reef, transition reef, and off reef) were defined to determine large-scale habitat use patterns. Throughout the area, at least 99 fish species were identified, many (19%) of which yielded new distributional data. Most species observed with the JSL were on prime reef (n ¼ 50) and transition reef (n ¼ 42) habitats, but the off reef habitat supported a well developed, but different fauna (n ¼ 25 species). Prime reef was characterized by Laemonema melanurum (21% of total), Nezumia sclerorhynchus (17% of total), Beryx decadactylus (14% of total), and Helicolenus dactylopterus (10% of total). The off reef areas were dominated by Fenestraja plutonia (19% of total), Laemonema barbatulum (18% of total), Myxine glutinosa (8% of total), and Chlorophthalmus agassizi (7% of total). Transition habitat exhibited a mixture of species that were also found on either prime reef or off reef habitats. Nezumia sclerorhynchus was the most abundant (25% of total) transition habitat species, followed by L. barbatulum (16% of total) and L. melanurum (14% of total). Several species (e.g., Anthias woodsi, B. decadactylus, Conger oceanicus, and Dysommina rugosa) demonstrated specificity to deep-reef habitats, while others (e.g., C. agassizi, Benthobatis marcida, F. plutonia, and Phycis chesteri) were always more common away from reefs. In addition to new distributional data, we provide behavioral and biological observations for dominant species. r 2007 Elsevier Ltd. All rights reserved.
Keywords: Deep-sea corals; Southeastern USA; Blake Plateau; Deep-reef fishes; Lophelia
1. Introduction ÃCorresponding author. Tel.: +1 910 395 3905; fax: +1 910 790 2292. Author's personalDeep-sea or cold-water copy corals form important E-mail address: [email protected] (S.W. Ross). habitat along many continental slopes and are 1Currently assigned (through Intergovernmental Personnel Act) to: US Geological Survey, Center for Coastal & Watershed receiving increasing attention worldwide. There is Studies, St. Petersburg, FL, USA. growing appreciation of their functions as fish
0967-0637/$ - see front matter r 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.dsr.2007.03.010 ARTICLE IN PRESS 976 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 habitat (Costello et al., 2005; Stone, 2006), as influence the distribution or survival of an organ- repositories of data on past environmental condi- ism, it is generally impossible to quantify all of tions (Adkins et al., 1998; Williams et al., 2006), and these. For this paper we concentrated on the as focal points of biodiversity. Deep coral habitats influence of deep-sea coral or rock (reef) structures are much more extensive than previously known, on the composition of benthic slope fish assem- and they face a number of threats (Rogers, 1999; blages. We considered the reefs to be mesoscale Koslow et al., 2000; Morgan et al., 2006; Roberts features (similar scale to hydrothermal vents or food et al., 2006). These high-profile features may falls) falling between megascale deep-sea structures, concentrate exploitable resources and enhance local such as mid-ocean ridges, canyons, or seamounts, productivity in ways similar to seamounts (Rogers, and microscale structures such as crevices, burrows, 1994; Koslow, 1997). Also, like seamounts, the or ripples. unique, diverse habitats occurring along the con- Basic habitat affinities have been established for tinental slope of the southeastern United States shallow-water (o200 m) reef fishes because such (SEUS) have escaped detailed examination not only habitats lend themselves readily to direct observa- because of their depths, but because the bottom tion methods that allow faunal classifications based topography is rugged and overlain by strong on habitat features. Stark (1968) defined levels of currents (i.e., Gulf Stream). Deep coral reefs habitat affinity (i.e., primary, secondary) for reef populate the SEUS continental slope in great fishes, which has facilitated general sea-floor habitat abundance (Stetson et al., 1962; Paull et al., 2000; classifications (Miller and Richards, 1980; SEA- Popenoe and Manheim, 2001; Reed, 2002a; Reed MAP-SA, 2001). Choat and Bellwood (1991) and Ross, 2005; Reed et al., 2006). By one estimate defined ‘‘reef fish’’ based on an array of character- (Hain and Corcoran, 2004), the SEUS and Gulf of istics (e.g., group features, ecological, habitat, Mexico have the most extensive deep coral popula- distributional, taxonomic, body morphology). In tions in US waters; however, the deeper parts of the deep-sea, however, conclusions about fish and these large regions are poorly explored. habitat relationships are likely biased by the indirect Habitat association data for deep-sea fishes are sampling (e.g., dredges, trawls) used to collect most lacking, leading to a general assumption that data. These methods confound and integrate fish/ habitat selection is opportunistic. Deep-sea fishes habitat associations over the distances sampled, and have often been grouped by depth (e.g., Jacob et al., in many cases the habitat sampled is unknown. 1998; Bull et al., 2001); however, even this basic Long term, replicated, direct observation is the habitat parameter may not accurately or strictly preferred methodology for gathering explicit data classify fish assemblages (Snelgrove and Haedrich, on faunal relationships to habitat in marine 1985; Haedrich and Merrett, 1990; Chave and environments (Starr et al., 1995; Connell et al., Mundy, 1994). Compared with continental shelf 1998; Cailliet et al., 1999). Such methods are and estuarine systems, deep-sea bottoms on the rarely applied in the deep-sea, and when used, scale of 100s to 1000s of meters are more homo- results ranged from only qualitative fish/habitat geneous, soft substrata. There are large, rugged descriptions (e.g., Wenner and Barans, 2001; Reed structures (canyons, mid-ocean ridges, trenches, et al., 2006) to detailed habitat association data seamounts) in the deep-sea, but much of the habitat indicating that several fishes were correlated with diversity that exists there, particularly deeper than rocky, reef habitats (Pearcy et al., 1989; Stein et al., 1000 m, may result from biological constructs (e.g., 1992; Chave and Mundy, 1994; Yoklavich et al., tubes, mudballs) that have ecological impacts (e.g., 2000). increased species richness) at a micro scale (Gage Despite increasing research on deep coral sys- and Tyler, 1991). In contrast, there are abundant tems, quantitative data on fishes associated with cases for close, often obligate, fish associations with these habitats are relatively lacking. In the cool various shallow-waterAuthor's marine and estuarine habitats personaltemperate to boreal northeasterncopy Atlantic Morten- (e.g., corals/hardgrounds: Parker and Ross, 1986; sen et al. (1995), Husebø et al. (2002), and Costello Lough et al., 1989; Stein et al., 1992; Chittaro, 2004; et al. (2005) noted that reefs formed by Lophelia Quattrini and Ross, 2006; vegetation: Adams, 1976; pertusa seemed to be important to some fishes. Lubbers et al., 1990; other: Pederson and Peterson, However, Auster (2005) suggested that in the 2002). While any of the biological or physical/ northwestern Atlantic deep corals were no more chemical parameters composing a habitat may important to fishes than other reef structures. ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 977
A similar conclusion of habitat opportunism was 2. Materials and methods reached for fishes associated with corals and sponges at moderate depths off southern California 2.1. Study areas (Tissot et al., 2006). Deep coral ecosystem data for fishes from the SEUS and Gulf of Mexico are Some deep coral study areas in the SEUS region limited, with studies reporting only a few taxa, have been named (e.g., Reed and Ross, 2005; Reed many not identified to species, from only a few et al., 2005, 2006), giving the impression that coral locations (Messing et al., 1990; Wenner and Barans, habitats are disjunct. Coral habitats on the Blake 2001; Reed et al., 2005, 2006). Data presented here Plateau (North Carolina to Cape Canaveral, FL) represent the first extensive, quantitative treatment are generally larger and more continuous than these of fishes on deep coral and hardground slope names imply. Detailed mapping of the region habitats of this region. Our overall goals were to combined with ground truthing are needed to clarify document species occurrences and relative abun- coral habitat distributions and the extent to which dances and to describe the degree of general habitat areas require discrete names. specificity of fishes on and around deep reef habitats Based on literature and our surveys, we desig- on the SEUS slope. By documenting fish distribu- nated eight general research areas where deep corals tions and abundances over reef and non-reef occurred in order to conduct replicate sampling habitats, we determined the degree to which fishes over several years (Fig. 1). During this study were specifically and predictably associated with (2000–2006) we steadily expanded our sampling to these continental slope habitats. include larger areas of the SEUS slope. Deep coral
Author's personal copy
Fig. 1. Deep-sea coral habitat study areas along the southeastern United States slope sampled during this study, 2000–2006. ARTICLE IN PRESS 978 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 habitats within the eight study areas vary in bottom sponges on tops of mounds/ridges, which are topography, coral development, and depth; how- surrounded by sand and rubble. ever, all of these reefs occur under the Gulf Stream and experience strong bottom currents. 2.2. Collection methods Off North Carolina, large mounds and ridges have been formed, which appear to be a sediment/ During eight (2000–2006) summer–fall cruises, coral rubble matrix topped with almost monotypic benthic and benthopelagic fishes on and near deep- stands (up to 3 m high) of live and dead L. pertusa. water coral banks were documented along the The three North Carolina Lophelia banks (desig- SEUS slope at depths of 356–910 m (Table 1, nated as Cape Lookout A, Cape Lookout B, and Fig. 1). Benthic sampling was conducted at eight Cape Fear, Fig. 1) are the northernmost deep-sea general localities (Fig. 1, see above) along the slope coral banks off the SEUS and the shallowest of our from central North Carolina to central Florida study sites (Table 1). The banks are generally similar using the Johnson-Sea-Link (JSL) submersible in physical attributes, rising about 80–100 m above (Harbor Branch Oceanographic Inst.), supplemen- the seafloor over o1 km distance and exhibiting ted with otter trawls. slopes up to 601. Extensive coral rubble zones The JSL was the primary gear used to collect data surround the mounds for large distances, and in on fishes and associated habitats. Usually two places seem to be quite thick (several cm at least). daytime dives (ranging from ca. 1.5–3 h each) were South of Cape Fear, deep coral habitat is more made with the JSL each day on or near coral banks variable and diverse and occurs deeper than off (Table 1). Our goal was to survey a variety of North Carolina. Sediment/coral mounds (Popenoe habitat types during each dive; however, since our and Manheim, 2001) are topped with a variety of priority was the largely unexplored coral mounds, anthozoans. The abundant hard bottoms (often more time was spent in that habitat. Scientists in exhibiting high-profile) on the Blake Plateau also both bow and stern compartments made frequent provide substantial substrata for L. pertusa and observations and operated hand-held digital, color other anthozoans. The Stetson area, a large region video cameras (Sony TRV 950) and digital audio of extremely rugged topography and diverse bottom recorders. In addition, an external, bow-mounted types (Stetson et al., 1962; Reed, 2002a; Reed et al., digital color video camera (Sony 3-chip CCD 2006), supports a variety and often a high density of camera with Canon 6–48 mm zoom lens) recorded corals (e.g., L. pertusa, Enallopsammia profunda, continuously throughout most dives. Two Leiopathes spp., Keratoisis spp.) and sponges. The (2001–2005) or four (2000) laser pointers (25 cm Savannah area is composed of numerous mounds apart) mounted on the camera were used to estimate and ridges of varying topography, and appears to fish lengths and measure habitat characteristics. have a heavier sediment load compared with other Depth, temperature, salinity, date, and time were sites. The bottom is covered with mostly dead logged at p1 scan s 1 intervals (data were overlain Lophelia, both rubble and standing thickets, with on the external camera video tapes) using a real- scattered low-profile living corals (e.g., L. pertusa, time data logger (Sea-Bird SBE 25 or 19plus) Stylaster spp.) and sponges (Reed, 2002a; Reed et attached to the submersible. Digital still images al., 2006; pers. obs.). Many habitats in the were taken with an externally mounted Nikon 955 Jacksonville area, especially on the northern end, camera (3.34 megapixel CCD, 4 8–115 mm zoom are composed of large, high-profile rock ledges with lens) at irregular intervals during dives in varying quantities and types of attached corals. 2003–2005. During all dives, the submersible’s Bottom types in this area are diverse, and some position was tracked from the surface support ship sampling sites are composed of Lophelia mounds using a Trackpoint II system. with mixed soft corals and sponges. Topographic We attempted to employ video transects standar- highs, most with coralAuthor's development, are abundant personaldized by time or distance copy (Parker and Ross, 1986; and nearly continuous from the Jacksonville area to Sulak and Ross, 1996); however, highly variable and south of Cape Canaveral (Ayers and Pilkey, 1981; rugged bottom topography coupled with often Paull et al., 2000; Reed, 2002a; Reed et al., 2006). strong currents made this impractical. Video re- Our southernmost study sites, divided into North corded when the JSL was stationary and moving Cape Canaveral and South Cape Canaveral, are (‘‘transecting’’) was used for both qualitative and composed of L. pertusa with mixed soft corals and quantitative fish and habitat assessments. When ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 979 C) Salinity 1 Temp ( Depth Range (m) 48.33 400–402 47.22 386–420 8.1 (5.5–9.0) 35.1 (34.9–35.3) 47.25 385–470 8.6 (5.6–10.6) 35.2 (34.0–35.8) 46.21 384–447 9.6 (6.3–10.9) 35.3 (33.9–36.0) 46.19 470–488 47.56 366–433 9.0 (8.4–9.6) 35.2 (35.1–35.4) 47.29 388–418 8.6 (6.2–9.4) 35.2 (34.1–35.6) 47.20 381–416 9.2 (9.0–10.1) 35.2 (34.7–35.7) 47.12 442–445 48.00 430–441 47.94 397–398 47.25 382–418 10.9 (8.9–12.0) 35.4 (34.0–36.1) 47.36 399–424 53.74 410–428 10.0 (9.6–10.4) 35.3 (35.1–35.4) 47.53 409–434 47.55 367–416 10.1 (9.8–10.5) 35.3 (35.0–35.5) 47.99 378–403 53.37 437–450 9.8 (9.6–10.2) 35.3 (35.0–35.4) 47.84 390–420 47.25 384–415 6.3 (5.9–6.9) 35.1 (34.9–35.2) 47.98 378–408 52.50 468–478 46.26 467–474 47.24 382–432 7.1 (6.2–8.3) 35.1 (34.9–35.2) 47.53 381–427 9.5 (9.1–9.9) 35.2 (35.0–35.6) 53.75 412–450 5.8 (5.4–6.0) 35.0 (35.0–35.1) 46.48 445–460 47.21 381–424 8.3 (7.5–9.0) 35.1 (34.8–35.3) 47.51 367–399 10.1 (9.2–10.6) 35.3 (34.9–35.7) 47.22 380–426 9.8 (9.5–10.0) 35.3 (35.2–35.4) 47.50 370–417 10.5 (9.9–10.8) 35.3 (35.1–35.5) 47.15 392–431 9.1 (8.0–9.8) 35.2 (35.0–35.3) 47.13 398–443 10.1 (9.0–10.9) 35.3 (35.0–35.5) 47.90 370–407 49.27 393–413 51.21 356–374 48.39 396–405 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 W) 1 End Longitude ( 18.99 75 19.47 75 19.45 75 19.51 75 19.37 75 19.42 75 19.48 75 19.10 75 17.58 75 19.66 75 19.45 75 19.65 75 11.41 75 18.52 75 19.50 75 19.56 75 10.77 75 18.93 75 19.40 75 19.38 75 12.27 75 18.79 75 19.42 75 19.69 75 11.42 75 19.31 75 19.48 75 19.42 75 18.36 75 19.44 75 19.41 75 19.51 75 18.77 75 19.34 75 18.49 75 17.28 75 19.08 75 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 N) 1 End Latitude ( 48.19 34 47.09 34 46.33 34 47.04 34 47.44 34 47.13 34 47.20 34 46.83 34 47.88 34 48.05 34 47.2 34 47.69 34 53.80 34 47.84 34 47.45 34 48.16 34 53.51 34 48.03 34 47.33 34 48.19 34 52.17 34 46.42 34 47.04 34 47.37 34 54.03 34 46.71 34 47.16 34 47.49 34 46.64 34 47.17 34 47.45 34 47.14 34 47.01 34 48.08 34 48.12 34 50.34 34 48.04 34 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 W) 1 Start Longitude ( 19.05 75 19.59 75 19.63 75 19.71 75 19.49 75 19.57 75 19.46 75 19.94 75 18.35 75 20.14 75 19.4 75 20.57 75 11.34 75 19.46 75 19.48 75 20.60 75 10.75 75 19.53 75 19.37 75 20.55 75 12.61 75 19.64 75 19.52 75 19.68 75 11.15 75 19.93 75 19.43 75 19.43 75 19.12 75 19.43 75 19.42 75 19.44 75 18.84 75 20.06 75 18.69 75 17.65 75 19.13 75 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 N) 1 Start Latitude ( Time (min)
Author's personal copy Table 1 2000–2006 Johnson-Sea-Link (JSL) submersible (S) and otter trawlStation (T) station data at deep-water coral sites off the southeastern US Gear Date Time Total SJ-02-037 T 10-Aug-02 N 39 34 Cape Lookout A JSLI-4206 S 28-Jul-00 M 114 34 JSLII-3304 S 11-Aug-02 M 148 34 JSLI-4891 S 17-Oct-05 A 115 34 JSLI-4207 S 28-Jul-00 A 109 34 JSLII-3305 S 11-Aug-02 A 149 34 CH-01-110 T 29-Aug-01 M 29 34 CH-06-044 T 21-Sep-06 M 30 34 CH-01-092 T 28-Aug-01 M 30 34 JSLII-3306 S 12-Aug-02 M 147 34 CH-01-111 T 29-Aug-01 M 30 34 Cape Lookout B JSLI-4365 S 24-Sep-01 M 153 34 CH-01-094 T 28-Aug-01 M 45 34 JSLII-3307 S 12-Aug-02 A 47 34 CH-01-112 T 29-Aug-01 A 30 34 JSLI-4366 S 24-Sep-01 A 74 34 CH-01-096 T 28-Aug-01 M 45 34 JSLII-3430 S 23-Aug-03 A 155 34 CH-01-113 T 29-Aug-01 A 30 34 SJII-01-053 T 24-Sep-01 A 30 34 CH-01-097 T 28-Aug-01 A 45 34 JSLII-3431 S 24-Aug-03 M 136 34 JSLI-4361 S 22-Sep-01 M 159 34 JSLII-3429 S 23-Aug-03 M 136 34 CH-01-100 T 28-Aug-01 A 45 34 JSLII-3432 S 24-Aug-03 A 130 34 JSLI-4362 S 22-Sep-01 A 135 34 CH-01-109 T 29-Aug-01 M 29 34 JSLI-4692 S 15-Jun-04 M 124 34 JSLI-4363 S 23-Sep-01 M 165 34 JSLI-4693 S 15-Jun-04 A 127 34 JSLI-4364 S 23-Sep-01 A 171 34 SJ-04-025 T 15-Jun-04 A 30 34 SJ-02-034 T 10-Aug-02 M 30 34 JSLI-4890 S 17-Oct-05 M 127 34 SJ-02-035 T 10-Aug-02 A 30 34 SJ-02-036 T 10-Aug-02 N 33 34 ARTICLE IN PRESS 980 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 C) Salinity 1 Temp ( Depth Range (m) 54.09 0–431 27.67 389–402 9.1 (9.0–9.5) 35.1 (35.1–35.3) 53.85 411–419 51.90 419–425 27.77 394–411 11.7 (11.2–12.1) 35.5 (35.4–35.7) 53.14 423–443 52.19 419–430 53.79 387–440 10.5 (9.4–11.2) 35.4 (35.2–35.5) 27.77 372–399 8.1 (7.9–8.3) 35.1 (35.0–35.1) 52.13 406–440 53.04 408–455 53.74 407–442 9.9 (9.7–11.3) 35.3 (35.0–35.8) 27.95 404–443 8.0 (7.8–8.2) 35.1 (35.0–35.1) 53.15 430–438 54.25 415–431 44.54 657–910 40.49 592–622 10.9 (10.8–11.0) 35.4 53.15 458–465 50.98 500–504 52.33 370–411 8.8 (8.6–9.7) 35.1 (35.0–35.3) 40.93 624–640 9.9 (9.8–10.0) 35.2 53.69 461–469 27.90 369–449 9.1 (8.4–9.5) 35.2 (34.8–35.4) 52.28 366–420 9.1 (8.4–9.6) 35.2 (35.0–35.3) 36.20 666–672 12.2 (12.1–12.3) 35.5 (35.5–35.6) 27.87 369–394 9.2 (9.4–9.7) 35.2 (35.1–35.3) 53.36 397–450 7.5 (6.3–8.2) 35.1 (35.0–35.2) 27.91 368–431 10.2 (9.3–11.2) 35.3 (35.0–35.6) 53.02 390–413 7.8 (7.6–7.9) 35.0 (35.0–35.1) 27.70 380–434 8.7 (7.9–9.8) 35.1 (34.8–35.3) 53.02 395–411 27.89 368–397 9.1 (8.7–9.4) 35.2 (35.1–35.2) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 W) 1 End Longitude ( 10.74 75 34.36 76 11.99 75 14.28 75 34.59 76 11.54 75 13.43 75 11.28 75 34.17 76 13.13 75 11.95 75 11.41 75 34.65 76 10.68 75 11.08 75 12.94 75 02.01 77 10.43 75 10.68 75 14.08 75 02.04 77 09.48 75 34.43 76 14.19 75 49.15 77 34.46 76 11.00 75 34.33 76 12.96 75 34.48 76 13.42 75 34.44 76 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 N) 1 End Latitude ( 53.56 34 27.71 33 53.27 34 51.55 34 27.83 33 52.83 34 51.85 34 53.62 34 27.89 33 52.11 34 52.76 34 53.65 34 27.98 33 52.76 34 53.82 34 46.29 34 40.44 32 52.74 34 50.53 34 52.44 34 40.71 32 53.33 34 29.05 33 52.30 34 36.77 31 27.93 33 53.59 34 27.91 33 53.09 34 27.75 33 52.47 34 27.95 33 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 W) 1 Start Longitude ( 11.65 75 34.37 76 15.28 75 34.57 76 12.37 75 14.40 75 11.28 75 34.18 76 14.00 75 12.78 75 11.41 75 34.64 76 12.33 75 11.88 75 12.56 75 01.75 77 11.37 75 11.44 75 13.90 75 02.01 77 10.36 75 34.33 76 14.00 75 49.15 77 34.38 76 10.66 75 34.38 76 12.96 75 34.28 76 14.23 75 34.38 76 12.94 75 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 N) 1 Start Latitude ( Time (min) 19-Sep-06 N 14 34 Author's personal copy ) a continued Table 1 ( Station Gear Date Time Total CH-06-026 T 20-Sep-06 M 30 34 JSLI-4697 S 17-Jun-04 A 120 33 CH-06-015 T 19-Sep-06 M 30 34 CH-06-027 T 20-Sep-06 M 31 34 JSLI-4694 S 16-Jun-04 M 132 34 JSLI-4896 S 20-Oct-05 M 146 33 CH-06-016 T 19-Sep-06 A 30 34 CH-06-028 T 20-Sep-06 M 30 34 JSLI-4695 S 16-Jun-04 A 130 34 JSLI-4897 S 20-Oct-05 A 125 33 CH-06-017 T 19-Sep-06 A 30 34 CH-06-029 T 20-Sep-06 A 30 34 SJ-04-035 T 16-Jun-04 A 30 34 Stetson JSLII-3419 S 17-Aug-03 M 131 32 CH-06-018 T 19-Sep-06 A 30 34 CH-06-031 T 20-Sep-06 A 30 34 JSLI-4892 S 18-Oct-05 M 140 34 JSLII-3420 S 17-Aug-03 A 126 32 CH-06-019 T 19-Sep-06 A 30 34 Cape Fear JSLII-3308 S 13-Aug-02 M 149 33 JSLI-4893 S 18-Oct-05 A 121 34 JSLI-4689 S 13-Jun-04 M 120 31 CH-06-021 T JSLII-3425 S 21-Aug-03 M 146 33 JSLI-4894 S 19-Oct-05 M 157 34 JSLII-3426 S 21-Aug-03 A 147 33 JSLI-4895 S 19-Oct-05 A 149 34 JSLII-3427 S 22-Aug-03 M 138 33 CH-06-012 T 19-Sep-06 M 33 34 JSLII-3428 S 22-Aug-03 A 126 33 CH-06-013 T 19-Sep-06 M 30 34 JSLI-4696 S 17-Jun-04 M 114 33 ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 981 depth ranges were afternoon (1200–1900 h EDT), ¼ 36.79 660–703 11.0 (10.9–11.8) 35.4 (35.3–35.5) 36.77 658–721 11.0 (10.9–11.1) 35.4 (35.3–35.5) 12.10 505–558 7.7 (7.5–7.8) 35.0 37.31 709–783 6.7 (6.7–6.9) 34.9 28.47 549–646 8.0 (7.2–8.7) 35.1 (35.0–35.2) 11.56 507–543 7.4 (7.4–7.6) 35.0 37.24 760–773 6.8 (6.8–7.0) 34.9 (34.8–35.0) 29.02 540–603 8.6 (8.1–9.2) 35.1 (35.0–35.3) 39.72 543–581 10.5 (10.3–10.9) 35.3 (35.1–35.5) 37.38 712–738 6.6 (6.5–6.7) 34.9 (34.9–35.0) 40.16 613–633 7.6 (7.3–8.0) 35.2 (35.1–35.2) 39.62 548–571 9.6 (9.0–10.5) 35.2 (34.9–35.5) 36.96 741–755 6.7 (6.7–6.8) 34.9 36.74 649–705 9.7 (8.2–11.8) 35.3 (34.9–35.8) 36.12 802–809 36.75 735–745 6.3 (6.3–6.4) 34.9 05.66 497–541 9.1 (9.0–9.1) 35.1 37.93 626–652 7.8 (7.8–8.0) 35.0 (34.9–35.0) 36.78 679–725 6.3 34.9 11.59 505–532 8.2 (8.2–8.3) 35.0 39.18 591–638 9.9 (9.8–10.0) 35.2 (35.2–35.3) 40.25 549–560 05.53 500–544 9.2 (9.1–9.3) 35.1 (35.1–35.2) 39.60 558–567 7.6 (7.5–8.3) 35.0 (34.9–35.1) 07.39 507–508 38.38 645–674 7.4 (7.3–7.5) 34.9 (34.9–35.0) 07.31 497–519 8.1 (8.1–8.2) 35.0 (35.0–35.1) 38.50 517–553 7.9 (7.5–8.3) 35.0 (34.9–35.1) 39.41 568–628 7.3 (7.2–7.6) 35.0 (34.9–35.0) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 morning (0600–1200 h EDT), A ¼ 49.56 77 50.75 77 46.43 79 47.60 79 16.17 77 46.62 79 47.75 79 15.83 77 30.97 79 47.61 79 00.95 77 30.84 79 46.62 79 50.79 77 29.86 79 02.53 79 44.52 79 48.70 79 02.38 79 46.56 79 30.10 79 05.46 79 44.57 79 30.85 79 42.30 79 28.93 79 42.32 79 48.03 79 31.26 79 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 36.69 31 36.72 31 12.26 31 37.19 28 28.42 32 11.64 31 37.30 28 28.82 32 39.62 30 37.40 28 40.00 32 39.62 30 36.96 28 36.83 31 36.14 30 36.82 28 06.09 31 37.81 30 36.84 28 11.70 31 39.09 30 39.89 28 06.16 31 39.68 30 07.42 31 38.50 30 07.88 31 38.39 30 39.63 30 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 409–431 m). 49.45 77 50.89 77 46.91 79 47.55 79 15.94 77 46.49 79 47.76 79 15.84 77 31.05 79 47.70 79 01.12 77 30.94 79 46.62 79 50.81 77 30.29 79 02.64 79 44.36 79 48.81 79 02.16 79 46.45 79 30.13 79 05.24 79 44.36 79 30.76 79 42.36 79 28.94 79 42.26 79 48.15 79 31.12 79 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0–409 m) and on the bottom (
Author's personal copy night (1900–0600 h EDT). Tucker trawl towed throughout the water column ( ¼ a JSLI-4698 S 18-Jun-04 M 109 31 JSLI-4699 S 18-Jun-04 A 130 31 JSLI-4898 S 21-Oct-05 M 118 32 JSLI-4906 S 30-Oct-05 A 86JSLI-4682 31 S 9-Jun-04 A 122 28 JSLI-4899 S 21-Oct-05 A 123 32 Jacksonville JSLI-4683 S 10-Jun-04 M 143JSLI-4702 S 30 20-Jun-04 M 124 28 JSLI-4903 S 27-Oct-05 M 111 32 JSLI-4684 S 10-Jun-04 A 126JSLI-4703 S 30 20-Jun-04 A 104 28 JSLI-4904 S 27-Oct-05 A 143 31 SJ-04-009 T 10-Jun-04 A 30South Cape Canaveral JSLI-4704 30 S 21-Jun-04 M 124 28 Savannah JSLI-4687 S 12-Jun-04 M 101 31 JSLI-4685 S 11-Jun-04 M 135JSLI-4705 30 S 21-Jun-04 A 110 28 JSLI-4688 S 12-Jun-04 A 93 31 JSLI-4686 S 11-Jun-04 A 113SJ-04-043 30 T 21-Jun-04 A 30 28 JSLI-4900 S 22-Oct-05 A 134 31 JSLI-4700 S 19-Jun-04 M 90Data are 30 only fordetermined the from Seabird period data when loggersN (at the stations JSL where or no ranges trawl are was listed, on there was the no bottom. variability For in data). JSL M stations, mean temperatures (and ranges), mean salinities (and ranges), and JSLI-4901 S 23-Oct-05 M 9 31 JSLI-4701 S 19-Jun-04 A 99 30 JSLI-4902 S 26-Oct-05 A 139 31 JSLI-4907 S 1-Nov-05 M 139 30 JSLI-4905 S 30-Oct-05 A 143 31 JSLI-4908 S 1-Nov-05 A 137 30 North Cape Canaveral JSLI-4681 S 9-Jun-04 M 122 28 ARTICLE IN PRESS 982 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 moving, we standardized operations as much as specifically, a main objective was to determine possible by keeping the JSL as close to the bottom whether there is a primary (possibly obligate) reef as practical, maintaining slow speed, tilting the fish fauna on deep coral habitat; therefore, broad external camera somewhat downward (toward generic habitat classification was appropriate. Vi- the seafloor), and videotaping on wide-angle view. deotapes and scientist observations were used to The consistent camera field of view and motion of develop the three general habitat classifications: the JSL across all transects helped standardize the prime reef, transition reef, and off reef (Fig. 2). fish counts among transects across different habi- Prime reef habitat had variable profile (X1m) tats. Each ‘‘transect’’ could have multiple video and usually occurred on or near tops of mounds segments (see below) used for statistical analyses. (Fig. 2a–c). Prime reef generally had dense coverage When the JSL stopped, the camera was variously set of corals (e.g., scleractinians, stylasterines, alcyona- from wide-angle to close-up views depending on ceans, gorgonians, and antipatharians) and/or subject matter and objectives. During the frequent sponges (e.g., hexactinellids, desmosponges). A high stops, specimens were first videotaped and then (450%) percentage of live coral frequently oc- collected using a suction device or grab, often curred in prime reef habitat. Some prime reef areas, supplemented with rotenone. Collections with asso- however, consisted of high profile, standing bushes ciated video were critical in confirming species of dead coral (L. pertusa), while others consisted of identifications on video. When specimens could extensive, moderate to high-profile (X1 m) rock not be collected, we recorded as much high-quality ledges with or without attached fauna. Sand video as possible. Specimens were preserved at sea channels were often interspersed throughout prime in 10% formalin seawater solution and later reef habitat. Transition reef included areas of transferred to 40% isopropanol, identified and mostly dead coral rubble with occasional scattered measured. Most species were measured in mm live corals and/or sponges. Transition reef usually standard length (SL); myxinids, sharks, anguilli- had moderate to low (o1 m) profile and occurred forms and macrourids were measured in mm total on the faces of steep slopes, near bottoms of slopes, length (TL); batoids were measured in mm disk or on the tops of slopes adjacent to prime reef areas width (DW). (Fig. 2d–f). At times, small rock outcrops with low- Otter trawls were deployed around coral banks to profile (o1 m) were present in transition habitat. sample benthic fishes (Fig. 1). Prior to trawling, the Patches of soft sediment often occurred in this area was surveyed with single beam sonar to ensure habitat. Off reef habitat was on relatively flat that trawls avoided major coral areas; however, the bottom (Fig. 2g–i), and soft sediment with occa- trawling objective was to tow as close to coral mounds sional coral or rock rubble was the dominant as possible. During 2001–2002, 2004 and 2006 the substratum. otter trawl (4.9 m head rope, 38.1 mm mesh) was towed for 29–45 min at 2 knot (3.7 km/h) ground 2.4. Data analyses speed. A Tucker Trawl (4 m2 opening, 1.59 mm mesh), deployed for a mid-water sample, was accidentally External video camera recordings were the main towed on the bottom (409–431 m) in 2006 for 14 min data used to describe the fish community and at 2 knot ground speed. Since this produced an associated habitats on and around deep coral banks. obvious sample of the bottom (collection included Videos from each dive were split chronologically several bottom fishes and substantial amounts of into distinct time segments during which fishes were coral, brittle stars, urchins), data from this tow were identified and counted and habitats were classified included with the otter trawl data. Trawl catches were (see Section 2.3). Video segments were designated preserved, identified and measured (mm SL, TL, or when the JSL stopped or started movement, the DW) as described above. Voucher specimens were camera zoom changed, the video quality changed deposited in the NorthAuthor's Carolina Museum of Natural personal(e.g., ‘‘dead time’’), orcopy when the habitat changed. Sciences ichthyology collection. Depth was recorded at the beginning and end of each video segment. All video ‘‘dead time’’ was 2.3. Habitat definitions removed from the dataset; this occurred when fish data could not be recorded because video was To determine large-scale habitat use patterns, turned off, was out of focus or clouded by sediment, three general habitat types were designated. More the camera was too high off the bottom, or digital ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 983
Fig. 2. Examples of habitat types: (a) Conger oceanicus in a prime Lophelia pertusa reef off North Carolina—JSLII-3432, 385 m; (b) mixed corals in prime reef in Stetson area—JSLI-4699, 669 m; (c) Polyprion americanus near a rock ledge in Stetson area—JSLI-4904, 663 m; (d) transition reef in Stetson area—JSLI-4903, 620 m; (e) Laemonema melanurum in Stetson transition reef—JSLII-3419, 610 m; (f) bamboo coral and sponges in transition area off Jacksonville, Florida—JSLI-4907, 541 m; (g) off reef, soft substrate habitat off North Carolina— JSLI-4366, 441 m; (h) debris on bottom in North Carolina off reef habitat—JSLI-4366, 449 m; and (I) Breviraja claramaculata in North Carolina off reef—JSLII-3304, 406 m. still images were taken. When data were difficult to Habitat specificity of benthic fishes was determined extract from the external camera videotapes, the using wide-angle transect video data. This analysis internal bow videotapes aided in species identifica- was restricted to fishes positively identified to unique tions and habitat classifications. Specimens col- taxa, with overall abundances X3 individuals. A lected (and videotaped) with the JSL enabled us to smaller abundance cutoff would not allow for the visually distinguish various species, many of which possibility of a species occurring in all three-habitat have rarely or never been seen in situ. types. For each species, relative (%) abundances were Species composition and relative abundances calculated by dividing the number of individuals in a were compared among prime, transition, and off particular habitat by the total number of individuals reef habitat types. To compare abundances of all of the same species from all habitats. species within a particular habitat, relative (%) Multivariate analyses were used to determine abundances were calculated (number of individuals differences in benthic fish assemblages among reef per species per habitat/totalAuthor's number of individuals personalhabitat types. All analyses copy were conducted in observed per habitat 100) using wide-angle trans- PRIMER 6 and based on guidelines in Clarke and ect video segment data (wide-angle transect ob- Warwick (2001) and Clarke and Gorley (2006). servations represent a subset of all observations). First, (using only transect data) sample units were Species occurrences were noted for all other times, established as numbers of each species per habitat when the submersible was stopped or the camera category (prime, transition, or off reef) per dive; the was videotaping in close-up mode. few samples with no species present were removed ARTICLE IN PRESS 984 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 from the dataset. Second, species’ abundances were and Warwick, 2001). The Bray–Curtis similarity standardized per sample by dividing the number of coefficient was used to calculate similarities between individuals per species by the total number of fishes species across samples. The resulting similarities per sample. Standardization was used because were clustered using group average linking to transect times were variable. Standardized abun- construct a dendrogram. On the dendrogram, each dances were then fourth root transformed to down habitat was listed after each species in decreasing weight the abundant species relative to the rare order of usage by the species. species. Similarities between samples were then Otter trawl catch data supplemented video calculated using a Bray–Curtis similarity coefficient. observations, adding to the overall species composi- A non-metric multidimensional scaling ordination tion of deep coral bank areas. Since all trawls were (MDS) plot and a dendrogram with group average towed near the banks in areas of low relief, the linking were created based on the Bray–Curtis catches represented transition or off reef habitats. similarity matrix. Finally, a one-way analysis of Catches that included dead coral rubble, live coral similarities (ANOSIM) and post-hoc multiple com- colonies, sponges and/or hard substrata represented parison tests were used to determine whether there tows in transition reef habitat. All other tows were were significant differences among fish assemblages classified as off reef habitat. in different reef types. SIMPER exploratory analy- sis was used to determine which species contributed 3. Results to the dissimilarities among reef types. In addition, inverse hierarchical cluster analysis Sixty-five JSL dives were completed on deep coral (PRIMER 6) was used to group species with similar habitats from off Cape Lookout, NC to just south numerical distributions across samples. To mini- of Cape Canaveral, FL (Table 1, Fig. 1). The dives mize rare species confounding the cluster analysis resulted in 136 h of bottom time and 116 h of (Field et al., 1982; Clarke and Warwick, 2001), useable video time (Tables 1 and 2). In all eight species that were o1% abundant across all habitats locations, prime reef was observed during 60 dives were removed from the dataset. Numbers of fishes (366–770 m) and transition reef was observed during were then standardized per species (number of 64 dives (367–783 m). Off reef habitat was observed individuals per species per sample/the total number during 25 dives (390–783 m) in seven locations of individuals per species), which reduced disparities (Table 2); it was not observed in the Stetson area. in species counts yet maintained ratios of occur- Time spent among methods (transects and stops) rences between species (Field et al., 1982; Clarke and habitats varied (Table 2). The JSL completed
Table 2 Fishes observed using submersibles (2000–2005) across three general habitat types. Number of dives and depth ranges (in parentheses) are under habitat type
Taxa Prime Reef Transition Reef Off Reef 60 (366–770 m) 64 (367–783 m) 25 (390–783 m)
TOTOT O 14.8 h 50.7 h 16.1 h 29.8 h 2.6 h 2.7 h
Myxinidae Undetermined 0.27 X 0.11 X 0.27 X Eptatretus sp. (2, 175–202) 0.18 X X Eptatretus minor (1, 227) X Myxine glutinosa X 7.69 X Chimaeridae Author's personal copy Chimaera monstrosa 0.09 0.11 0.27 Squalidae Squalus spp. 0.09 0.11 X Cirrhigaleus asper (1, 785) 0.36 X 0.11 Squalus cubensis 0.27 X 1.35 X 0.27 X Odontaspididae Odontaspis ferox X ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 985
Table 2 (continued )
Taxa Prime Reef Transition Reef Off Reef 60 (366–770 m) 64 (367–783 m) 25 (390–783 m)
TOTOT O 14.8 h 50.7 h 16.1 h 29.8 h 2.6 h 2.7 h
Scyliorhinidae Scyliorhinus spp. 0.18 0.34 Scyliorhinus hesperius 0.18 Scyliorhinus meadi 0.27 X 0.45 X Scyliorhinus retifer (1, 440) 0.45 X 2.47 X 0.27 Carcharhinidae Carcharhinus altimus XX Narcinidae Benthobatis marcida (1, 172) 2.39 Rajidae Undetermined 0.22 2.39 X Breviraja claramaculata 0.53 Dactylobatus armatus X Dipturus cf. sp. X 0.27 Fenestraja plutonia (1, 98) 0.90 X 18.83 X Mobulidae Mobula hypostoma X Synaphobranchidae Dysommina rugosa (6, 130–215) 0.90 X Synaphobranchus spp. 1.81 X 5.62 X 5.31 X Synaphobranchus kaupii (2, 233–341) 0.09 X Congridae Undetermined X Conger oceanicus 7.78 X Nettastomatidae Nettenchelys exoria (5, 93–447) 0.27 X 0.22 X Argentinidae Argentina cf. striata 1.27 X 0.22 X 0.27 Sternoptychidae Maurolicus weitzmani (128, 35–45) 7.87 X 11.36 X 2.92 X Polyipnus clarus (5, 46–56) 0.45 X 0.34 X 12.20 X Sternoptyx sp. X Stomiidae Chauliodus sloani X Ateleopodidae Undetermined 0.11 Chlorophthalmidae Chlorophthalmus agassizi (2, 100–125) 1.12 X 6.63 X Paralepididae Undetermined X 0.11 0.27 Myctophidae Undetermined X 1.24 X Diaphus dumerilii (215, 17–63) X Bythitidae Bellottia apoda (1, 44) X Bythites gerdae (1, 71) X Bythites cf. gerdae 0.45 X Diplacanthopoma brachysomaAuthor's(1, 221) personal X copy Macrouridae Undetermined 0.45 2.12 Nezumia spp. 0.22 X 0.80 X Nezumia aequalis 0.22 X 0.27 X Nezumia cf. bairdii 0.09 X Nezumia sclerorhynchus (6, 145–228) 17.01 X 25.08 X 4.24 X ARTICLE IN PRESS 986 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007
Table 2 (continued )
Taxa Prime Reef Transition Reef Off Reef 60 (366–770 m) 64 (367–783 m) 25 (390–783 m)
TOTOT O 14.8 h 50.7 h 16.1 h 29.8 h 2.6 h 2.7 h
Moridae Laemonema spp. 1.00 X 1.46 X 0.80 Laemonema barbatulum (10, 63–158) 3.53 X 15.97 X 17.51 X Laemonema melanurum (1, 220) 20.81 X 14.40 X 0.80 Physiculus cf. fulvus 0.09 X X Physiculus karrerae (1, 250) 0.09 X Phycidae Phycis chesteri 0.22 X 2.39 Urophycis cf. chuss 0.18 Merlucciidae Merluccius albidus 0.34 X 3.45 X Lophiidae Lophiodes beroe (6, 280–400) 0.36 X 1.12 X Lophiodes monodi (1, 325) X Lophius gastrophysus (1, 210) 0.09 0.11 Chaunacidae Chaunax stigmaeus (6, 95–255) 0.67 Ogcocephalidae Dibranchus atlanticus 0.22 X Trachichthyidae Hoplostethus occidentalis (11, 91–164) 5.61 X 1.01 X Berycidae Beryx decadactylus 13.94 X Zeidae Zenopsis conchifera X Scorpaenidae Undetermined 0.45 X 0.56 X 0.80 X Helicolenus dactylopterus (8, 66–252) 10.32 X 7.87 X 5.57 X Idiastion kyphos (7, 65–109) 0.36 X 0.22 X Phenacoscorpius nebris (1, 55) X Pontinus rathbuni (2, 119–185) X X Setarches guentheri (2, 106–132) 0.09 0.22 0.27 X Trachyscorpia cristulata (3, 60–230) 1.27 X 2.02 X X Acropomatidae Synagrops sp. A X Synagrops sp. B 0.34 X 0.27 Polyprionidae Polyprion americanus 0.36 X 0.67 X Serranidae Anthiinae 0.18 X Anthias woodsi 0.81 X Hemanthias aureorubens X Epigonidae Undetermined X Echeneidae Undetermined X Trichiuridae UndeterminedAuthor's personal X X copy Xiphiidae Xiphias gladius X
Hours of observation spent in each habitat are listed by method. Species % relative abundances were calculated for each habitat during transects (T). Occurrences (denoted by X) during other (O) times are noted. Number of individuals collected via submersible, followed by length (mm) range, are indicated (in parentheses) after each species. ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 987
14.8 and 16.1 h of transects in prime and transition exhibited a mixture of species that could also be habitats, respectively, and 2.6 h of transects in off found on either prime reef or off reef. N. reef habitat. sclerorhynchus was the most abundant (25% of Salinities at deep-coral banks were stable across transition reef total) transition habitat species, study areas, while temperatures fluctuated among followed by L. barbatulum (16% of total) and L. dives within and across all study areas (Table 1). melanurum (14% of total). Over all dives and study areas, mean bottom We used multivariate analyses on the video data salinities ranged from 34.9 to 35.5 (70.0 SE). Mean to examine fish assemblage differences among bottom temperatures ranged from 5.8 1C(70.0 SE) habitat types. MDS and hierarchical clustering at a location within Cape Lookout B to 12.2 1C analyses were performed on 42 benthic fish species (70.0 SE) at a location within the Stetson area. observed across 142 samples in prime, transition, During a few dives, observers noted benthic and off reef habitat types. The ANOSIM test temperature fluctuations over short distances, where revealed weak, but significant (Global R ¼ 0.2, temperatures increased or decreased briefly, and p ¼ 0.1%), fish assemblage differences among the during these events, the observers saw mixing of three reef types; however, the post-hoc comparison density layers in the water column. tests indicated moderate difference between prime To supplement the JSL data, 33 otter trawl and off reef habitats (R ¼ 0.5) and no differences (includes the bottom Tucker trawl) tows were between off and transition habitats (R ¼ 0.2) and completed around deep coral banks in 356–910 m transition and prime habitats (R ¼ 0.1) (R ¼ 0 depth (Table 1). In the Cape Lookout A area when groups are the same and R ¼ 1 when groups (356–488 m), eleven tows were completed in transi- are different; Clarke and Warwick, 2001). The low tion reef, and five tows were in off reef. Ten tows Global R value was a result of the similarity were completed in off reef habitat, and four tows between transition and prime reef and transition were in transition reef habitat in the Cape Lookout and off reef habitat types. Since transition reef B area (395–504 m). One tow was in deep habitat is an ecotone between prime and off reef (657–910 m) off reef habitat east of the Cape habitats, the lack of significant fish assemblage Lookout B area. Farther south, only one tow was differences between transition reef and the other made off Jacksonville in transition habitat two habitat types was not surprising. (802–809 m), and one tow was completed off South To examine off and prime reef fish assemblage Cape Canaveral in off reef habitat (549–560 m). differences in greater detail, we reanalyzed the dataset (PRIMER 6, Section 2.4) using only fish 3.1. Submersible data counts from these two reef types (39 species, 78 samples). The resulting MDS plot (Fig. 3) displayed We identified at least 66 unique fish taxa in 38 families from the submersible data over all locations and years (Table 2). While most of the species richness was within prime reef or transition habitats (50 and 42 species, respectively), the soft substrata off reef habitats also supported a well developed, but different fish fauna (25 species) (Table 2). The ichthyofauna of all three general habitat types was dominated by relatively few species, with little overlap in species between prime reef and off reef habitats. In particular, prime reef was characterized by Laemonema melanurum (21% of prime reef total), Nezumia sclerorhynchusAuthor's(17% of total), personal copy Beryx decadactylus (14% of total), and Helicolenus dactylopterus (10% of total). The off reef areas were dominated by Fenestraja plutonia (19% of off reef Fig. 3. Multidimensional scaling (MDS) ordination of 78 off and prime reef samples based on the Bray-Curtis similarity matrix total), Laemonema barbatulum (18% of total), calculated from standardized, fourth root transformed fish Myxine glutinosa (8% of total) and Chlorophthal- abundances (39 species). Clusters are defined at a 30% (black mus agassizi (7% of total). Transition habitat outlines) similarity level. ARTICLE IN PRESS 988 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 a good representation (Stress ¼ 0.11) of the samples plutonia (off). SIMPER analysis also indicated low in two-dimensional space (Clarke and Warwick, average similarity of samples within each habitat 2001). The MDS indicated three off reef groupings (off reef samples: 32% similar, prime reef samples: (two groupings consisted of one sample each) and 33% similar). The relatively moderate statistical four prime reef groupings (one consisted of only one difference between off and prime reef habitats, sample) at a 30% level of similarity. The ANOSIM combined with the low average similarity of samples test revealed moderate, but significant, differences within a particular habitat, resulted from a location (Global R ¼ 0.5, p ¼ 0.1%) between the two (eight study sites along the SEUS) effect on the habitats, while SIMPER exploratory analysis in- dataset. Location differences will be discussed in a dicated that prime and off reef assemblages were future paper where data will be further analyzed by 88% dissimilar. Species driving the differences study site (Ross and Quattrini, manuscript in included the following abundant species: L. barba- preparation). tulum (off reef), L. melanurum (prime), N. scler- We further examined the habitat specificity of the orhynchus (prime), H. dactylopterus (prime) and F. abundant benthic species observed during transects
Author's personal copy
Fig. 4. Within species relative (%) abundances across three habitat types. n ¼ number of individuals observed during wide-angle transects. ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 989
Fig. 5. Dendrogram of fish species using group average linking from a Bray-Curtis species similarity matrix calculated from standardized abundances. The 19 most abundant benthic species were included in this analysis. Three major groups are 95% dissimilar, denoted by the dotted vertical line through the dendrogram. Letters (in parentheses) denote order of abundance in habitats, P ¼ prime reef, T ¼ transition reef, O ¼ off reef. by comparing the relative abundances within a abundant in prime and transition reefs clustered species across habitat types (Fig. 4). Five species together, and fishes most abundant in off reef areas were observed only in prime reef: Anthias woodsi, B. grouped together, except Synaphobranchus spp. that decadactylus, Bythites cf. gerdae, Conger oceanicus were most abundant in transition habitats clustered and Dysommina rugosa. Species that were most with the off reef group. The third major cluster abundant in prime reef habitat and were rarely consisted of only one species, Squalus cubensis, observed in off or transition reef areas included which was more abundant in transition reef than Hoplostethus occidentalis, Cirrhigaleus asper, L. other habitats and was seen most frequently in the melanurum, Nettenchelys exoria, and H. dactylop- Savannah study area. Within the prime/transition terus. Some species were more abundant in off reef reef species group, there were two subgroups at high habitat, and never occurred on prime reef, including (50%) similarity levels; one subgroup consisted of Benthobatis marcida, C. agassizi, Merluccius albidus, B. decadactylus with C. oceanicus, and another M. glutinosa, F. plutonia and Phycis chesteri. subgroup consisted of L. melanurum with N. Certain species were more abundant in transition sclerorhynchus. B. decadactylus and C. oceanicus reef compared with other habitats, including Chau- were abundant in prime reef habitats, especially off nax stigmaeus (observed only in this habitat), North Carolina, while L. melanurum and N. Lophiodes beroe, Scyliorhinus retifer and L. barba- sclerorhynchus were abundant in both prime and tulum. The large, commercially important wreckfish transition habitats in several deep coral bank areas. (Polyprion americanus) was also commonly ob- Within the off reef species group, M. albidus and P. served in transition habitat. These data clearly chesteri grouped together at a high ( 60%) illustrated (Fig. 4)Author's species-specific general habitat personalsimilarity level, and these copy two species were common use differences. only off North Carolina. Inverse cluster analysis also indicated habitat- While our analyses concentrated on benthic related groupings of fish species. The 19 most fishes, several species of pelagic and midwater fishes abundant species from 142 video samples clustered also visited these deep reefs. Three solitary Xiphias into three major groups that were 95% dissimilar gladius were observed briefly in prime reef coral from one another (Fig. 5). Fishes that were most habitat during dives at the Cape Lookout coral ARTICLE IN PRESS 990 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 banks. One individual swam rapidly up a steep reef Table 3 face, whereupon it struck the submersible and swam Numbers of fishes followed by length (mm) ranges (in parenth- away. Three very large Mobula hypostoma also eses) collected with the otter trawl in two habitats occupied the Cape Lookout coral banks during a Taxa Transition Reef Off Reef few dives. Individuals occurred singly and were n ¼ 16 n ¼ 17 observed circling above the crests of Lophelia (370–809 m) (356–910 m) mounds. An unidentifiable echeneid was attached Myxinidae on the dorsal head region of one M. hypostoma. Eptatretus sp. 1 (202) Additionally, several species of mesopelagic fishes, Myxine glutinosa 5 (250–320) 54 (186–345) including Diaphus dumerilii, Maurolicus weitzmani Etmopteridae and Polyipnus clarus, were observed in aggregations Etmopterus bullisi 2 (135–190) near the bottom in all three habitats (Table 2). Scyliorhinidae Scyliorhinus retifer 5 (123–330) 1 (183) Torpedinidae 3.2. Otter trawl data Torpedo nobiliana 1 (231) Rajidae We collected 69 fish species (including mesopela- Breviraja claramaculata 3 (124–171) 3 (45–128) gics) in transition and off reef habitats using the Dipturus teevani 2 (234–255) 1 (261) Fenestraja plutonia 26 (72–118) 69 (28–129) otter trawl (Table 3). Of these, 49 species were Synaphobranchidae caught in tows from transition reef habitat, and 49 Dysommina rugosa 21 (79–210) 2 (180–183) species were collected in off reef habitat tows. Synaphobranchus kaupii 1 (316) Trawling added 34 benthic species to the overall Ophichthidae composition of fishes on and around SEUS deep Ophichthus sp. 1 (292) Congridae coral banks and included several families that were Bathycongrus vicinalis 1 (115) not observed with the JSL (e.g., Draconettidae, Nettastomatidae Etmopteridae, Grammicolepidae, Ipnopidae, Para- Nettenchelys exoria 1 (345) lichthyidae and Peristediidae). Several species col- Serrivomeridae lected by trawl may have been observed with the Serrivomer sp. (damaged) 1 (148) Argentinidae JSL, but they could not be accurately identified on Argentina striata 1 (131) video. These included Dipturus teevani, Epigonus Gonostomatidae pandionis, Synagrops bellus, S. spinosus and S. Cyclothone spp. (damaged) 9 trispinosus. Cyclothone microdon 1 (44) Of the total number of fishes collected by trawl in Cyclothone pallida 3 (37–53) Sternoptychidae each habitat, H. dactylopterus was the most Maurolicus weitzmani 1 (37) abundant in both transition (44%) and off reef Polyipnus clarus 1 (23) (22%) habitats (Table 3). Also, L. barbatulum Sternoptyx diaphana 1 (32) (18%) and P. chesteri (17%) were abundant in Stomiidae transition and off reef habitats, respectively. Some Chauliodus sloani 3 (87–160) Ateleopodidae fishes that were common in trawl collections, such Ateleopus spp. 2 (451–520) 2 (340–365) as Dibranchus atlanticus, Setarches guentheri and Ijimaia antillarum 1 (208) Urophycis regia, were rarely observed with the JSL Chlorophthalmidae probably because of reduced observation time in off Chlorophthalmus agassizi 41 (51–121) 58 (49–129) reef habitats. Parasudis truculenta 1 (180) Ipnopidae Juveniles of several species whose adults were Bathypterois bigelowi 2 (127–146) commonly observed with the JSL in transition or Neoscopelidae prime reef habitats were collected with the otter Neoscopelus macrolepidotus 4 trawl in off reef habitatAuthor's (Table 3). These included personalMyctophidae copy Nezumia spp. (aequalis or sclerorhynchus, species Benthosema glaciale 1 (52) Diaphus dumerilii 10 (56–60) identification not resolvable because of damage or Polymixiidae missing scales), L. barbatulum, Trachyscorpia cris- Polymixia lowei 1 (153) tulata and H. occidentalis. Juvenile H. occidentalis Ophidiidae may have been collected in the water column Benthocometes robustus 1 (112) because in the otter trawl tow containing this Bythitidae Bythites gerdae 1 (52) ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 991
Table 3 (continued ) Table 3 (continued )
Taxa Transition Reef Off Reef Taxa Transition Reef Off Reef n ¼ 16 n ¼ 17 n ¼ 16 n ¼ 17 (370–809 m) (356–910 m) (370–809 m) (356–910 m)
Macrouridae Paralichthyidae Undetermined (damaged) 1 Hippoglossina oblonga 1 (180) Caelorinchus caelorhincus 1 (209) 6 (177–260) Cynoglossidae Hymenocephalus italicus 19 (105–170) Symphurus nebulosus 1 (63) Malacocephalus occidentalis 1 (362) 1 Total Number 884 870 Nezumia sp. (damaged) 1 Nezumia sp. (aequalis/ 3 22 (60–82) n ¼ number of tows, followed by sample depth ranges (in sclerorhynchus) parentheses). Damaged specimens could not be positively Nezumia bairdii 2 (260–299) identified to species and/or accurately measured. Nezumia sclerorhynchus 3 (94–160) 1 (107) Ventrifossa macropogon 1 (290) Moridae species, several mesopelagic species (e.g., Cyclothone Gadella imberbis 3 (114–147) 3 (102–134) spp., Sternoptyx diaphana) were also collected. Laemonema barbatulum 161 (62–188) 128 (54–192) Juvenile Hoplostethus mediterraneus were collected Physiculus karrerae 4 (130–190) Phycidae by otter trawl in off reef habitat; this species was not Phycis chesteri 12 (195–284) 147 (162–254) collected in any other habitat. Also, juveniles of Urophycis regia 61 (93–310) 13 (142–217) common off reef species were collected in non-reef Urophycis tenuis 1 (373) habitat, including newly hatched Breviraja clarama- Lotitdae culata (n ¼ 1, 45 mm DW) and F. plutonia (n ¼ 11, Enchelyopus cimbrius 1 (105) Merlucciidae 28–44 mm DW). Merluccius albidus 11 (132–388) 10 (112–241) Lophiidae 3.3. Behavioral observations Lophiodes beroe 1 (235) Lophius gastrophysus 4 (135–237) A unique advantage of using submersibles is the Chaunacidae Chaunax suttkusi 1 (28) 1 (121) ability to observe and record behaviors of fishes in Ogcocephalidae situ. Despite possible observational artifacts intro- Dibranchus atlanticus 34 (29–117) 53 (26–138) duced by the submersible (from noise and lighting), Trachichthyidae many fishes exhibit few reactions to the submersible, Hoplostethus mediterraneus 4 (22–50) and the direct observations seem worthwhile be- Hoplostethus occidentalis 3 (77–131) 2 (19–63) Grammicolepidae cause many of these fishes have not been observed Xenolepidichthys dalgleishi 1 (73) in the wild, and literature generally lacks data on Scorpaenidae behavioral attributes of deep-sea fishes. Costello Helicolenus dactylopterus 385 (30–335) 190 (50–220) et al. (2005) reported in situ behaviors of fishes Idiastion kyphos 1 (94) associated with coral banks, noting that 75% of Setarches guentheri 24 (34–144) 12 (51–129) Trachyscorpia cristulata 11 (65–160) 14 (47–156) species did not react to the underwater cameras. Peristediidae Other studies reported wide ranges of reactions to Peristedion ecuadorense 1 (140) 4 (42–153) submersibles or ROVs, but maintained that direct Peristedion greyae 5 (64–120) 2 (102–109) observations were valuable (Chave and Mundy, Peristedion truncatum 1 (48) 1 (68) 1994; Uiblein et al., 2002, 2003; Lorance and Acropomatidae Synagrops bellus 8 (15–146) 1 (148) Trenkel, 2006). Behavioral observations for the Synagrops spinosus 1 (106) 3 (63–133) dominant species (Fig. 4) are summarized below, Synagrops trispinosus 1 (73) except such data appear in Sections 3.4 and 3.5 for Epigonidae Author's personalfishes yielding new distributional copy information, and Epigonus pandionis 8 (105–145) 2 (109–121) lophiiform observations were discussed by Caruso Percophidae Bembrops gobioides 1 (152) et al. (2007). Draconettidae M. glutinosa was frequently observed with JSL Centrodraco acanthopoma 6 (80–97) 2 (83–90) and collected with the otter trawl in off reef habitat. Bothidae This species was never observed buried in soft Monolene sessilicauda 1 (40) sediments in this study, but it was always observed ARTICLE IN PRESS 992 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007
Fig. 6. In situ photographs of noteworthy species: (a) Eptatretus sp.—JSLII-3429, 415 m; (b) Chimaera monstrosa—JSLI-4905, 506 m; (c) Scyliorhinus hesperius—JSLI-4686, 604 m; (d) Scyliorhinus meadi—JSLII-3431, 409 m; (e) Odontaspis ferox—JSLI-4903, 627 m; (f) Carcharhinus altimus—JSLI-4365, 393 m; (g) Conger oceanicus (large eel) and Dysommina rugosa (small eel)—JSLI-4694, 387 m; and (h) Nettenchelys exoria—JSLI-4699, 717 m. swimming rapidly over the bottom. At a soft bottom on soft substrata, off reef habitats. At times substratum study site off Cape Hatteras it was one individuals buried into the sediments. Occasionally, of the most abundant fishes, often observed buried F. plutonia was observed lying on dead coral rubble with only the snout exposed (SWR, pers. obs.). in transition habitat. Most sharks were observed in reef areas and Species of Anguilliformes, a dominant group of exhibited various behaviors. Squalids were highly fishes, exhibited various behaviors on deep coral mobile throughout reef habitats, occurring 1–5 m banks. Synaphobranchus spp., nearly ubiquitous off the bottom. Several individuals, especially S. throughout habitats off Florida, were observed in cubensis, were usually in constant motion and at constant motion swimming 0.5–5 m above the times appeared to be attracted to the submersible. substrate. Synaphobranchus spp. occurred singly or Small S. cubensis were also observed resting on the in small groups (o10 individuals). We collected two bottom. Cirrhigaleus asper was usually mobile, but Synaphobranchus kaupii; but because species within some individuals were observed in close contact the genus Synaphobranchus cannot be visually with the bottom. S. retifer exhibited many beha- identified, we could not confirm whether all viors, including sittingAuthor's motionless on the bottom personalindividuals observed copy were this species. S. kaupii (mostly on coral rubble or dead coral matrix), exhibited a variety of locomotion behaviors (for- moving rapidly around coral mounds or the nearby ward movement, station holding, and drifting) that open substrate, and occasionally rubbing their varied with habitat (including depth, substrate type, bodies on dead Lophelia branches. and current speed) in the Bay of Biscay (Uiblein The batoid fishes, F. plutonia and B. marcida, et al., 2002, 2003). C. oceanicus adults ( 700–1000 mm were consistently observed lying motionless on the TL), never observed off-reef, were intimately ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 993
Fig. 7. In situ photographs of noteworthy species: (a) undetermined ateleopodid—JSLI-4894, 443 m; (b) Nezumia sclerorhynchus—JSLI- 3419, 613 m; (c) Helicolenus dactylopterus—JSLII-3431, 413 m; (d) Idiastion kyphos—JSLI-4693, 416 m; (e) Setarches guentheri- JSLI-4906, 542 m; (f) Trachyscorpia cristulata—JSLI-4687, 499 m; (g) Hoplostethus occidentalis within coral matrix—JSLII-3305, 383 m (h) Beryx decadactylus—JSLII-3431, 384 m; and (i) Anthias woodsi—JSLI-4362, 368 m. associated with the coral habitat. On a few These three species were observed resting on, occasions individuals were observed swimming hovering over, or slowly swimming above soft around the corals, but most observations were of substrata or coral rubble. On occasion, pairs of single individuals with either the heads or tails M. albidus and pairs of P. chesteri were observed. protruding from the coral matrix (Figs. 2a and 6g). An abundant fish of deep coral banks, N. This eel was rarely observed in rocky habitat and sclerorhynchus, was regularly observed south of was most abundant where corals (especially Lophe- Cape Fear, NC. Most individuals hovered o0.5 m lia) were the dominant substratum. D. rugosa was from the substrate with their heads angled down- observed several times in the vicinity of C. oceanicus ward toward the seafloor. Several individuals were (Fig. 6g). Several individuals, all relatively small also observed swimming over the open substrate, (collected individuals, 130–215 mm TL), were ob- and no individuals were observed within the coral served swimming around the heads of C. oceanicus. bushes. N. sclerorhynchus was easily identified in D. rugosa occurred only in prime reef habitat, and videos because of its small size, slender body, and was frequently observed with head in, tail out of tall, white first dorsal fin (Fig. 7b). high-profile LopheliaAuthor'sbushes. Occasionally, this eelpersonalL. melanurum was frequentlycopy and consistently was not seen until flushed out from coral rubble or observed around coral areas. Although its distribu- bushes using rotenone, indicating that they spend tion was highly skewed toward reef habitat, it was some time deeper within the coral matrix. observed over less dense reef cover as our surveys The mostly off reef species, C. agassizi, M. albidus expanded toward the southern Blake Plateau. A and P. chesteri, generally occurred as solitary wide range of sizes ( 150–350 mm TL) was individuals closely associated with the bottom. observed, and individuals were easily recognized ARTICLE IN PRESS 994 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 by the distinct black markings on the caudal and guentheri dramatically changes as the diagonal posterior anal and dorsal fins (Fig. 2e). They were bands disappear and the body becomes brick red usually moving slowly just above the bottom. On or orange, with dark reddish to black spots along its several occasions, L. melanurum were observed in body and a dark reddish to black stripe along its pairs, and generally, one individual was much ventral midline. This fresh dead coloration is smaller than the other. Its congener, L. barbatulum, commonly described in the literature (e.g., Poss overlapped L. melanurum in habitat usage but also and Eschmeyer, 2002). ranged further away from reefs, occurring over soft T. cristulata was common, especially on deep substrata. coral banks south of North Carolina (Fig. 7f). All B. decadactylus occurred occasionally as single individuals sat motionless on the bottom, also the individuals, but was more often observed in dominant behavior of the subspecies T. c. echinata aggregations ranging from o10 to 450 individuals. in the eastern Atlantic (Lorance and Trenkel, 2006). All were large adults that were usually swimming A few were observed resting on top of Lophelia over (1–3 m above) and around the reef (Fig. 7h); branches, but most were on coral rubble, either on however, a few individuals were seen under ledges open substrata or next to black corals, bamboo created by Lophelia bushes. B. decadactylus were corals, sponges, or Lophelia bushes. The coloration most common around the highest profile reefs of T. cristulata was generally red, with light, small (usually at the tops of mounds) with dense coral spots scattered on the body and reddish-brown bars coverage. B. decadactylus appears to be closely from the head to caudal peduncle. Additionally, associated with rugged, steep terrain; other studies pale to white blotches were apparent on the head, have also documented this same pattern of habitat and beneath the dorsal fin. usage (Chave and Mundy, 1994; Popenoe and P. americanus was the largest fish consistently Manheim, 2001). observed (Fig. 2c) in this study. Usually we Literature and our data indicate that H. dacty- observed these fish in loose aggregations of 3–420 lopterus uses a wide range of habitats. It was the individuals. They hovered just above the bottom, most common scorpaenid on the reefs, and adults sometimes touching it, or swam slowly 1–2 m above and juveniles were observed perched under coral the bottom. Individuals often exhibited no response bushes, perched on top of coral (live or dead), or to the submersible, or at times seemed distracted by sitting on the substrate near corals (Fig. 7c). This it and moved away. While only observed in reef species is also common on coral habitat at Rockall habitats, it tended to be most common around the Banks off Ireland where it displays the same bases and slopes of reef mounds. Polyprion amer- behavior as noted above (SWR, pers. obs.). When icanus was observed on tops of mounds only when observed away from reef habitat, it was nearly the coral was low profile and mostly dead, or where always closely associated with whatever structure rock ledges occurred (Stetson area). They were was available (burrows, anemones). This species is never observed in dense, high profile, live coral widely known from trawl samples over supposedly habitat. Several studies have documented P. amer- soft substrata (Haedrich and Merrett, 1988; Gordon icanus to be abundant in coral mound and high et al., 1996), but trawls obscure habitat data. relief rock habitats on the Charleston Bump (see Regardless of habitat details, H. dactylopterus is Sedberry, 2001). strongly associated with the bottom (Uiblein et al., 2003). 3.4. New fish distributional data for the SEUS slope S. guentheri, a common scorpaenid on open, soft substrata (S.W. Ross, unpubl. data), was rarely Nineteen fish species that we observed or observed during this study. When observed, indivi- collected yielded new depth and/or geographic duals were on soft substrata or on coral rubble, range data. Caruso et al. (2007) also documented sometimes sitting nextAuthor's to structures (e.g., Lophelia personalfour Lophiiform fishes copy collected during this study colony). Live coloration of S. guentheri is pale red, that were new to the region. with darker red, diagonal bands and markings along The genus Eptatretus may be more often asso- its body and a dark red to black spinous dorsal fin, ciated with coral (and/or structure) than previously tipped in pale red/white (Fig. 7e). This coloration thought (Fernholm and Quattrini, in press). Epta- seems more pronounced in smaller individuals. tretus minor was collected with the JSL in prime reef Upon reaching the surface, the coloration of S. habitat in the Stetson area in 561 m (NCSM 44650, ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 995
227 mm TL, JSLI-4899). We examined an addi- (Stehmann and Bu¨ rkel, 1984; Didier, 1998; identi- tional specimen of E. minor (GMBL 97-39) that was fication confirmed by D. Didier Dagit, pers. trapped on hard bottom habitat in 201 m off South commun.). Individuals were observed alone, hover- Carolina (D.M. Wyanski, pers. commun.), an ing over coral rubble, hard coral, or sandy unusually shallow depth for this genus in the substrate. Upon the submersible’s approach, indi- western North Atlantic. These records substantially viduals were disturbed and swam away. In the extend the geographic and depth ranges of E. minor. northeastern Atlantic, C. monstrosa was also Previously, this species was known from the north- reported from off reef (Mortensen et al., 1995; eastern Gulf of Mexico to the Dry Tortugas, FL in Freiwald et al., 2002; Costello et al., 2005) and coral 300–472 m (Fernholm and Hubbs, 1981; McEa- rubble habitats near Lophelia reefs (Costello et al., chran and Fechhelm, 1998). Fernholm and Hubbs 2005). (1981) reported that coral and mud were collected in An uncommon catshark, Scyliorhinus hesperius, two trawls that caught E. minor. They assumed that was observed twice off Jacksonville, FL resting on E. minor probably bury into sediment and thus, thick coral rubble in 580 m (JSLI-4908) and 604 m were caught in mud habitat. Two individuals of an (JSLI-4686) in prime reef habitat (Fig. 6c). Their undescribed species of Eptatretus (Fig. 6a)(Fern- sizes were estimated to be 500 mm TL. This holm and Quattrini, in press) were collected by JSL species was readily identified by unique coloration (UF 165853, JSLI-4364) in transition, rubble consisting of white spots scattered within black habitat in the Cape Lookout A site, and one saddles (identification confirmed by G.H. Burgess, individual was trawled (UF 165852, CH-06-017) in pers. commun.). These are the first records for the off reef habitat adjacent to the Cape Lookout B site. SEUS continental slope and the deepest yet for the In addition, 19 individuals of this new species were species. Previously, this species was known from observed with the JSL in prime and transition reef depths of 274–530 m on the continental slopes of the habitats, swimming over the bottom and/or moving western Caribbean Sea (Springer, 1966, 1979; through (or protruding from) the coral matrix Springer and Sadowsky, 1970; Compagno, 1984). (Fig. 6a). The unique bright pink color of this Six B. claramaculata were collected off Cape hagfish allowed us to easily distinguish it from other Lookout, NC by otter trawl (NCSM 44430, 132 mm hagfish species. DW, 409–434 m, CH-01-094; NCSM 44434, 124 We observed three, solitary adult Chimaera and 171 mm DW, 390–420 m, CH-01-096; NCSM monstrosa (all 600 mm TL) in prime, transition, 44436, 117 and 128 mm DW, 445–460 m, CH-01- and off reef habitats from the Savannah 100; NCSM 44649, 45 mm DW, 500–504 m, CH-06- (506–528 m, JSLI-4905, 4906) and North Cape 031). In addition, two adults were observed by JSL Canaveral (770 m, JSLI-4682) study areas (Fig. 2i) in off reef habitat resting on the bottom off (Fig. 6b). In addition, we examined a specimen of Cape Lookout, NC (406–469 m, JSLI-4206, JSLII- this species collected by trawl in the Tongue of the 3304). Previously, B. claramaculata was known Ocean, Bahamas in 1483 m (UF 166362, 880 mm from South Carolina to the Florida Keys in TL). This specimen appears to be the one reported 293–896 m (McEachran and Matheson, 1985). An by Sulak (1982). These represent the first records for additional individual was collected off North the western Atlantic. C. monstrosa was previously Carolina in 640 m (UF 29868). known from the eastern Atlantic in depths of A single ophichthid eel was collected by trawl 300–1171 m from off Greenland, Norway, the adjacent to the Cape Lookout B coral bank (CAS North Sea, and Iceland to the Azores, including 224335, CH-06-016). This eel represents a new the Mediterranean Sea (Stehmann and Bu¨ rkel, species of Ophichthus (McCosker and Ross, 1984; Moller et al., 2004; Mytilineou et al., 2005). in press). It may occur in shallower areas (40–100 m) during One Bathycongrus vicinalis was collected off Cape summer in the northernAuthor's part of its range (Stehmann personalLookout, NC from a copy transition reef area with a and Bu¨ rkel, 1984). Although we did not collect C. Tucker trawl (NCSM 44646, 115 mm TL, monstrosa, we recorded high quality video and 409–431 m, CH-06-021). This juvenile specimen identified this species using the following characters: possessed its larval pigmentation, three lateral rows anal fin separate from ventral caudal fin, silver color of melanophores (D.G. Smith, pers. commun.). The with brown spots and undulating stripes, grayish Tucker trawl was towed both in midwater and on fins, and black distal margins of unpaired fins the bottom (409–431 m), but most likely this species ARTICLE IN PRESS 996 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 was collected on the bottom. B. vicinalis is known One ateleopodid (visual identification not possible from the eastern Gulf of Mexico, northwestern beyond family level) was observed (Fig. 7a) with the Bahamas, West Indies, and Central and South JSL off Cape Lookout, NC swimming close to the America from 101 to 503 m (Smith and Kanazawa, bottom on the edge of Lophelia-sand habitat (443 m, 1977; Smith, 1989a; McEachran and Fechhelm, JSLI-4894). Our records extend the geographic 1998). ranges of these genera. N. exoria was observed and collected frequently One Benthocometes robustus was collected by from deep coral banks along the SEUS (Fig. 6h). otter trawl from transition reef habitat in the Cape Off North Carolina, one individual was collected Lookout B area (NCSM 44743, 112 mm SL, by otter trawl (NCSM 44467, 345 mm TL, 423–443 m, CH-06-015). This species was previously 396–405 m, SJ-02-036), and five were collected by reported in the eastern Atlantic off northwest Africa JSL (NCSM 44444, 143 mm TL, 374 m, JSLI-4363; and in the Mediterranean (Nielsen et al., 1999). In NCSM 44448, 190 mm TL, 425 m, JSLI-4365; NCSM the western Atlantic, B. robustus was reported from 44450, 447 mm TL, 440 m, JSLI-4366; NCSM 44454, off Cuba (Goode and Bean, 1895), northern Straits 107 mm TL, 414 m, JSLI-4695; NCSM 44455, 93 mm of Florida (Staiger, 1970), and along the northeast TL, 402 m, JSLI-4696). Also, an adult was observed US coast from Norfolk Canyon (USNM 326148) to by JSL off North Carolina (442 m, JSLI-4366); five Hudson Canyon (Musick et al., 1992; Moore et al., adults (all 300 mm TL) were observed in the 2003). This species was collected in bottom depths Stetson area (641–717 m, JSLI-4689, 4699, 4898); of 200–1652 m (Moore et al., 2003). Nielsen and two adults were observed off Jacksonville, FL Evseenko (1989) proposed that the disjunct dis- (614–631 m, JSLI-4686); and two were observed tribution of B. robustus could result from an off Cape Canaveral, FL (687–748 m, JSLI-4703, extended pelagic larval phase. While possible, it 4705). N. exoria was often observed with head in, seems likely that the distribution of this rarely tail out of the Lophelia coral matrix, but a few collected fish will prove to be more continuous on individuals were observed in the open, swimming both sides of the Atlantic. near the bottom. This poorly documented species Bellottia apoda was collected with the JSL off was previously reported off Cuba (Claro et al., Jacksonville, FL in 629 m (NCSM 44451, 44 mm 2000), the northeast coast of Florida and the SL, JSLI-4685) in prime, rocky reef habitat, Bahamas in 227–494 m (Smith et al., 1981; Smith, extending both its geographic and depth ranges. 1989b). This species was flushed out of a crevice in a rock Ateleopodidae is a poorly known family in need ledge with rotenone. B. apoda was previously known of revision (Smith, 1986; Moore, 2002a) whose from the subtropical eastern and western Atlantic, species have a circumtropical distribution (Proko- the Mediterranean Sea, the Galapagos Archipelago fiev, 2006) and have been infrequently collected in and the Philippines, occurring at bottom depths of 200–800 m (Moore, 2002a). Ijimaia antillarum was 30–569 m (Nielsen et al., 1999; Mytilineou et al., collected by otter trawl off Cape Lookout, NC 2005). (NCSM 44438, 208 mm SL, 378–403 m, CH-01- Four Physiculus karrerae individuals were col- 112). This species has been reported from Cuba, the lected by otter trawl (NCSM 44429, 130 mm SL, Gulf of Mexico (Howell Rivero, 1935; McEachran 409–434 m, CH-01-094; NCSM 44433, 175 and and Fechhelm, 1998) and New England (Moore 190 mm SL, 390–420 m, CH-01-096; NCSM 44643, et al., 2003). Species of the genus Ateleopus have 137 mm SL, 423–443 m, CH-06-015). In addition, been reported from the eastern Atlantic (e.g., Smith, one was collected and observed with the JSL 1986), Pacific (e.g., Prokofiev, 2006), and Indian (NCSM 44442 250 mm SL, 383 m, JSLI-4361), and oceans (e.g., Smith, 1986), and in the western three other adults were observed only with the JSL Atlantic, Ateleopus spp. have been reported from (384–424 m, JSLI-4361, 4365, JSLII-3429). All were off Suriname (Shimizu,Author's 1983) and the Gulf personal of observed in prime, Lopheliacopyreef habitats under Mexico (Moore, 2002a). Four individuals of coral bushes or dead coral matrices. Upon the Ateleopus spp. (not assignable to species) were submersible’s attempts to collect specimens with collected by otter trawl off Cape Lookout, NC rotenone, individuals swam out from the coral and (NCSM 44431, 520 mm SL, CH-01-096; NCSM appeared to quickly seek shelter under another coral 44437, 451 mm SL, 399–424 m, CH-01-111; NCSM bush/matrix. P. karrerae is known from a few 44465, 340 and 365 mm SL, 356–374 m, SJ-02-035). records off Bermuda (Smith-Vaniz et al., 1999), the ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 997
Gulf of Mexico (Sulak et al., in press), the also have been this species. Previously, this Caribbean, Brazil, St. Helena (Paulin, 1989), species was known from the Caribbean to Brazil Tristan da Cunha (Andrew et al., 1995) and the (Mochizuki and Sano, 1984; Cervigo´ n, 1993; Mejı´ a Walvis Ridge (Trunov, 1991) in 240–800 m (Smith- et al., 2001; Lopes et al., 2003) and the northeastern Vaniz et al., 1999). Gulf of Mexico (Ruiz-Carus et al., 2004) from Off Cape Lookout, NC six Idiastion kyphos depths of 137–550 m (Mochizuki and Sano, 1984; specimens were collected with the JSL (NCSM Ruiz-Carus et al., 2004). Additional trawl collec- 44443, 95 mm SL, 398 m, JSLI-4361; NCSM 44445, tions (MCZ 62079, 160714, 166112, 166167) exist 82 mm SL, 374 m, JSLI-4363; NCSM 44460, 65 mm from off Cape Hatteras, NC in 66–127 m. SL, 397 m, JSLII-3305; NCSM 44462, 107 mm SL, A. woodsi was observed (14 adults, 300 mm TL) 387 m, JSLII-3306; NCSM 44452, 81 mm SL, 416 m, with the JSL (Fig. 7i) off Cape Lookout, NC JSLI-4693; NCSM 44457, 90 mm SL, 443 m, JSLI- (367–407 m, JSLI-4362, 4363, 4891, 4892, 4893, 4894), and another was collected by a bottom 4895, JSLII-3306) in prime, Lophelia reef habitats. Tucker trawl tow (NCSM 44647, 94 mm SL, Individuals were solitary, closely associated with 409–431, CH-06-021). An additional specimen was reef habitat, and several were observed in or darting collected in the Stetson area (NCSM 44458, 109 mm into the Lophelia matrix. This species was pre- SL, 569 m, JSLI-4899). Ten other individuals (all viously known from collections off South Carolina, 80–100 mm TL) were observed with the JSL off the east coast of Florida, and the Dry Tortugas North Carolina (385–443 m, JSLI-4206, 4365, 4693, (Anderson and Heemstra, 1980) in 175–475 m 4697, 4890, JSLII-3304, 3306, 3430, 3431, 3432); (Heemstra et al., 2002). one ( 90 mm TL) was observed in the Stetson area Ten E. pandionis were collected by otter trawl off (560 m, JSLI-4899), and another ( 90 mm TL) was Cape Lookout, NC (NCSM 44428, 145 mm SL, observed in the Savannah study area (519 m, JSLI- 409–434 m, CH-01-094; NCSM 44435, 109 mm SL, 4905). Several I. kyphos were observed perched on 445–460 m, CH-01-100; NCSM 44439, 378–403 m, branches of live or dead L. pertusa. Some indivi- 113 and 121 mm SL, CH-01-112; NCSM 44441, 108 duals were sitting well within, while others were on and 117 mm SL, 378–408 m, CH-01-113; NCSM the fringes of, low to high relief coral bushes. All sat 44466, 105, 117 and 118 mm SL, 396–405 m, SJ-02- motionless and were not disturbed by the submer- 036; NCSM 44645, 121 mm SL, 406–440 m, CH-06- sible. All I. kyphos were pale red, but some 016). This amphi-Atlantic species was recorded individuals had large white blotches covering the in the western Atlantic from the southern New posterior portion of the head to under the spiny England slope (Moore et al., 2003), New Jersey, dorsal fin (Fig. 7d). When brought to the surface, Straits of Florida, the Gulf of Mexico and the blotches disappeared. I. kyphos was previously the Caribbean to French Guiana in 210–751 m known from scattered records in the Atlantic (Poss (Staiger, 1970; Mayer, 1974). Our records are the and Eschmeyer, 2002): off Venezuela (Eschmeyer, first documented for the SEUS south of Cape 1965), northeastern Florida and the eastern South Hatteras. Atlantic (Anderson et al., 1975) in depths of Eight Centrodraco acanthopoma individuals were 229–622 m. collected by otter trawl off Cape Lookout, NC Phenacoscorpius nebris was collected with the (NCSM 44427, 83 mm SL, 409–434 m, CH-01-094; JSL as it was sitting motionless on outer branches NCSM 44432, 81 mm SL, 390–420 m, CH-01-096; of a Lophelia bush in the Savannah study area in NCSM 44469, 97 mm SL, 370–407 m, SJ-04-025; 499 m (NCSM 44459, 55 mm SL, JSLI-4902). NCSM 44642, 80, 83, and 85 mm SL, 423–443 m, This collection extends both its geographic and CH-06-015; NCSM 44644, 83 and 90 mm SL, depth ranges. This species was reported in the 406–440 m, CH-06-016). This species occurs in both tropical eastern Atlantic (Mandritsa, 1992)in the eastern and western Atlantic (Fricke, 1992). In 320 m and in theAuthor's western Atlantic (Venezuela personalthe western Atlantic, copy adults are known from and the Gulf of Mexico) in 347–475 m (Eschmeyer, Georgia to the Florida Keys (Briggs and Berry, 1965, 1969). 1959) and the Gulf of Mexico (McEachran and Synagrops trispinosus was collected by otter trawl Fechhelm, 2005) in depths of 384–594 m (Briggs and off Cape Lookout, NC (NCSM 44464, 73 mm SL, Berry, 1959; Davis, 1966). Several pelagic juveniles 356–374 m, SJ-02-035). A few specimens (Synagrops have also been reported off New England (Moore spp.) that we observed with the JSL (Table 2) may et al., 2003). ARTICLE IN PRESS 998 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007
3.5. Noteworthy records margin, second dorsal fin smaller than first, and coloration. This deep-water carcharhinid has a Several poorly known species, rarely recorded in circumglobal distribution; however, records of it SEUS waters, were documented during this study. are sporadic (Compagno, 1984, 2002). In the Odontaspis ferox is a deep-water shark infrequently western Atlantic, C. altimus was recorded from reported throughout most of the world’s oceans near the surface (at night over deep-water) to 430 m (Compagno, 1984; Bonfil, 1995) in 13–420 m (Com- (Compagno, 1984; Anderson and Stevens, 1996) off pagno, 2002). In the western Atlantic, it is known Florida, the Bahamas, Cuba, the Atlantic coast of from the Yucata´ n shelf (Bonfil, 1995), the northern Mexico, the Yucata´ n, Costa Rica, Nicaragua, Gulf of Mexico (Sulak et al., in press), Cuba (Claro Venezuela, Brazil, the northeastern Gulf of Mexico and Parenti, 2001), Brazil (Compagno, 2002) and (Compagno, 1984; McEachran and Fechhelm, 1998; off Cape Hatteras, NC (Sheehan, 1998). We add Compagno, 2002) and off Cape Hatteras, NC records of two individuals of O. ferox to continental (Stillwell and Casey, 1976). US waters, both observed swimming slowly near the Four specimens of Bythites gerdae were reported bottom around the JSL: one individual ( 2000 mm from off Cape Hatteras, NC in 204–562 m (Nielsen TL, Fig. 6e) in the Stetson area (627 m, JSLI-4903), and Cohen, 2002), resulting from a related study and one ( 2500 mm TL) off Jacksonville, FL (S.W. Ross et al., unpubl. data). B. gerdae is also (573 m, JSLI-4683). O. ferox was visually distin- known from a few records in the Florida Straits in guished by the presence of five gill slits all anterior 786–832 m (Nielsen and Cohen, 1973). We add to the pectoral fin, high dorsal fins (second slightly records for this species off the SEUS: one individual smaller than the first), the first dorsal originating collected by otter trawl in transition reef habitat off well in advance of pelvic fin, a markedly high anal Cape Lookout, NC (NCSM 44426, 52 mm SL, fin placed well behind second dorsal, and coloration 409–434 m, CH-01-094), and one collected by JSL (identification confirmed by G.H. Burgess, pers. within a crevice in a Lophelia mound in the South commun.). Twelve individuals (all 600 mm TL) of Cape Canaveral study area (NCSM 44456, 71 mm the poorly documented Scyliorhinus meadi (Fig. 6d) SL, 687 m, JSLI-4705). Also, a few undetermined were observed off North Carolina (374–442 m, bythitids, possibly B. gerdae, were observed fre- JSLI-4207, 4361, 4364, 4693, 4892, 4893, JSLII- quently in prime, Lophelia habitat off North 3428, 3431, 3432) and one ( 600 mm TL) was Carolina. Individuals were observed protruding observed in the Savannah study area (498 m, JSLI- from or darting into large coral bushes. 4687). This species was visually identified by unique One Ventrifossa macropogon was collected by coloration of conspicuous dark saddles without otter trawl off Cape Lookout, NC (NCSM 44440, light or dark spots. Individuals were usually solitary 290 mm TL, 378–403 m, CH-01-112). V. macropo- and were generally observed swimming close to the gon was previously known from North Carolina bottom, often contacting or rubbing their bodies on (two records), the east coast of Florida, the the corals. They were also observed resting on the Caribbean to Guyana, and the Gulf of Mexico in bottom usually near coral formations (Fig. 6d). Few 439–1000 m (Marshall, 1973; Cohen et al., 1990). records of S. meadi have been reported in the Adult H. occidentalis (all 100–200 mm TL) were western Atlantic from off North Carolina, Florida, commonly observed (n ¼ 130) with the JSL off the Bahamas to Cuba, and the Yucata´ n in the Gulf North Carolina and Jacksonville, FL. Eleven other of Mexico in 329–549 m (Springer, 1966, 1979; individuals were collected using the JSL from Springer and Sadowsky, 1970; Burgess et al., 1979; reef areas off North Carolina (NCSM 44446, Compagno, 2002). Four Carcharhinus altimus (all 116 mm SL, 372 m, JSLI-4363; NCSM 44447, 120, 2000 mm TL) were observed with the JSL (Fig. 6f) 135, 137, 140, 155, and 164 mm SL, 400 m, JSLI- swimming rapidly near the bottom off North 4364; NCSM 44449, 151 mm SL, 423 m, JSLI-4365; Carolina (380–424Author's m, JSLI-4365, 4890, JSLII-3307, personalNCSM 44461, 135 mmcopy SL, 382 m, JSLII-3305; 3427). Visual identification (confirmed by G.H. NCSM 44463, 98 mm SL, 427 m, JSLII-3427; Burgess, pers. commun.) was based on a combina- NCSM 44453, 91 mm SL, 394 m, JSLI-4693). Five tion of characters: posterior gill slits located above more individuals were collected off North Carolina pectoral fin origin, midlength of first dorsal fin base by otter trawl, two of which were juveniles (NCSM closer to pectoral fin than pelvic fin insertion, first 44468, 110 mm SL, 396–405 m, SJ-02-036; NCSM dorsal fin moderately high with a straight anterior 44470, 19 and 63 mm SL, 657–910 m, SJ-04-035; ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 999
NCSM 44648, 77 and 131 mm TL, 415–431 m, (2006) listed Lophius sp. from one location. We CH-06-029). This species was thought to be rare never saw or collected Raja sp., which they listed along the coast of the SEUS (Moore, 2002b), with from three areas, but instead we observed and previous records from off New England (Moore collected several other Rajidae taxa throughout the et al., 2003), Bermuda (Smith-Vaniz et al., 1999), region. They indicated that Nezumia sp. might be North Carolina (MCZ 75974,137247), the east coast three species one of which was N. atlanticus; our of Florida, Florida Straits, the northern Gulf of data suggested N. sclerorhynchus to be more likely. Mexico, the Caribbean, southern Brazil, Venezuela, Reed et al. (2006) did not report abundances or the Guianas (Woods and Sonoda, 1973; Kotlyar, habitat associations, so no further comparisons are 1986), Cuba (Claro et al., 2000) and Suriname possible. George (2002) reported 24 fish species (13 (Uyeno and Sato, 1983) in 124–823 m (McEachran of which we report here) trawled from a possible and Fechhelm, 1998; Smith-Vaniz et al., 1999). H. coral area north of the Stetson sites. Recent occidentalis is closely associated with coral habitat multibeam sonar mapping of that area (S.W. Ross, and was one of the most cryptic fishes documented unpubl. data), however, suggests a soft substrate on coral habitats. Single individuals were usually habitat with no elevation. observed hidden deep within coral branches The Charleston Bump area (roughly between our (Fig. 7g) and were rarely observed swimming over Savannah and Stetson sites) contains rugged hard the reef substrate. We were unable to identify bottom with various coral coverage (Sedberry, species-specific colorations or morphologies to 2001). Weaver and Sedberry (2001) listed 15 fish visually distinguish the two Hoplostethus spp. taxa from the Charleston Bump, and only six of (H. mediterraneus and H. occidentalis) known from those coincided with species from our surveys. SEUS waters. However, since we only collected Differences in habitat (less coral and more rock at H. occidentalis with JSL, we recorded all observed Charleston Bump) and sampling methods could individuals as this species. Their cryptic behavior account for the fish biota differences between their and association with rugged habitats, indicate they study and ours. Popenoe and Manheim (2001) may be more abundant than our data suggest and observed that P. americanus, Beryx spp., and also may explain why captures were previously Trachichthyidae used large undercut ledges as lacking in the region. shelter in the Charleston Bump area. Undercut ledges of the size noted by Popenoe and Manheim 4. Discussion (2001) were not encountered in our surveys; yet, we also observed B. decadactylus and H. occidentalis We documented a diverse ichthyofauna (99 under ledges created by Lophelia bushes and lower species) on and near deep coral habitats of the profile rock ledges, suggesting a general tendency SEUS slope between 356 and 910 m. This represents for these species to seek shelter. the most fish species yet recorded from deep reef/ Two adjacent deep coral sites (in 308–536 m) were coral ecosystems in the Atlantic. There are few other surveyed in the north-central Gulf of Mexico (Sulak deep coral (4200 m) ecosystem references in the et al., in press). Most of the 36 fish taxa positively SEUS region with which to compare our fish data, identified to species from JSL video occurred on reef and those are generally qualitative (fishes neither habitat (mixed rock, corals and sponges). A few collected nor counted). All 12 of the fishes visually species (Hyperoglyphe perciformis, Gephyroberyx identified to species level by Reed et al. (2006) from darwini, Epinephelus niveatus, H. occidentalis, Gram- three study areas (Stetson, Savannah, Jacksonville) micolepis brachiusculus, C. oceanicus) were seen overlapping with our sites were also reported in our almost exclusively on reef habitat (Sulak et al., in study. Their identification of fewer fishes could press). Although all of the above species are known result from lack of voucher specimen collections from the SEUS slope, only two (H. occidentalis, (precluding some speciesAuthor's level identifications), diffi- personalC. oceanicus) occurred incopy our study primarily in reef culty of visual identifications for many species habitat. The lack of E. niveatus on SEUS slope reefs (many never before seen in situ), and lack of was likely due to greater depths sampled compared literature documentation for the region’s fishes to the Gulf of Mexico sites where this species was (many new records for the region). We found seen only around 310 m. Differences in data Lophiodes (two species) to be the most common treatment preclude further habitat related compar- lophiid (Caruso et al., 2007), while Reed et al. isons between these studies. Reed et al. (2006) ARTICLE IN PRESS 1000 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 reported nine fish taxa from coral and hard bottom depths) 37–48 km southwest of the Cape Fear deep habitats in the western Gulf of Mexico (412–558 m), coral reef (Quattrini and Ross, 2006), and only one all of which we also observed or collected on SEUS species (C. oceanicus) was shared between shelf edge sites. and slope reefs. Reed et al. (2006) noted that 73 fish Compared with other deep regions of the western species occurred on the Oculina Bank off east- Atlantic (Haedrich and Merrett, 1988; Diaz et al., central Florida (70–100 m), none of which were 1994; Sulak and Ross, 1996; Moore et al., 2003; shared with coral/hardground systems deeper than Powell et al., 2003), the slope environment of the 300 m. These examples typify the disparity between Blake Plateau has been poorly sampled for benthic the shelf and slope reef faunas in terms of species and benthopelagic macrofauna. This area has a richness and composition. Differences in tempera- disproportionate amount of high-profile rugged ture (usually 4151 on outer shelf vs. p12 1Con substrata, which has probably hampered extensive slope) probably play the biggest role in maintaining sampling via traditional means (dredges, trawls). the faunal separation of the two different SEUS reef Most of the species documented from off reef and to fish communities (shelf, subtropical versus slope some extent transition reef habitats were species species). known from the area, and were those we expected to In contrast to many shallow water fish species, the find. In contrast, many of the prime and transition lack of habitat association data for deep-water reef species were either previously unknown to the fishes has hampered our understanding of deep reef area (19% of total) or were thought to be rare (or communities and the roles of complex habitats in both). The apparent abundance and widespread structuring or maintaining deep-sea communities. occurrence of many of these fishes are further While fish associations with deep corals may be indications of close association with reef habitat, more opportunistic in some areas (Husebø et al., which seems to have secluded them from discovery. 2002; Auster, 2005; Costello et al., 2005; Stone, Outer continental shelf hardgrounds in the SEUS 2006), our direct observation data, spanning several are also difficult to sample, and, as for the deep years and a wide geographic area, suggest a stronger coral reefs, there were many additions to the fish tie to reef habitats along the SEUS slope. Our data, fauna when these areas were better sampled and those of Sulak et al. (in press), support the (Quattrini et al., 2004; Quattrini and Ross, 2006). hypothesis that deep, slope reefs function much like In comparison, none of the 23 fish species recorded shallow coral reefs, hosting a unique, probably from cold-water coral reef habitats from off Nor- obligate, ichthyofauna and concentrating food way to the Porcupine Seabight were new to the resources. Although not quantified, we observed region (Costello et al., 2005), probably because abundant small invertebrates (amphipods, krill, these reefs and nearby habitats have been histori- shrimps), a potential forage base, more frequently cally better sampled. Also, if northeastern Atlantic around dense coral habitat than over non-reef areas. slope fishes were habitat generalists, they would Costello et al. (2005) reported similar observations. have greater opportunity for their ranges to be well Whether corals themselves or only the structure documented (i.e., not secluded in hard to sample made by the corals (with its related benefits) are the habitats). attracting factors is unclear (Auster, 2005); never- The fish fauna of Blake Plateau deep reefs is theless, corals are a major contributor to deep-sea completely different from that of the relatively habitat complexity and structure (Roberts et al., better studied SEUS shelf hardgrounds. The shal- 2006). Comparisons of mobile macrofauna at slope lower (p200 m) continental shelf reef systems of the depths between rocky habitat with no or few SEUS consist of one Oculina reef off central Florida attached invertebrates (corals, sponges) and habitat (Reed, 2002b) and extensive emergent hardgrounds with dense coral cover would be instructive for from Cape Hatteras to southern Florida (SEA- evaluating the role of deep-sea corals in attracting MAP-SA, 2001).Author's These hardground communities personalnon-sessile fauna. copy are subtropical and are best developed along the Unfortunately, most data on deep-sea fishes shelf edge (Grimes et al., 1982; Barans and Henry, result from trawl or other indirect sampling, with 1984; Parker and Ross, 1986; Quattrini and Ross, little specific information on habitat. Several species 2006). Using similar methodology as in this study, important on Blake Plateau deep coral habitat (e.g., we documented 113 fish species on several southern Synaphobranchus spp., H. occidentalis, B. decadac- North Carolina shelf edge hardgrounds (57–128 m tylus, H. dactylopterus) have been captured ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 1001 elsewhere, but details of habitat were missing and systems. Although limited, data on the ages of often obscured by sampling techniques (e.g., Hae- western Atlantic deep coral systems indicate they drich and Merrett, 1988; Merrett et al., 1991; have provided habitat for tens of thousands of years Massuti et al., 2001; Moore et al., 2003). Some of (Ayers and Pilkey, 1981; Paull et al., 2000), thus the previous investigations of deep reef fishes do not allowing substantial time for a deep reef fauna to help elucidate the extent of the habitat use patterns evolve close habitat associations. we report because studies were either restricted to a An important function of most shallow complex few species, were geographically or temporally structure habitats (whether reef, vegetation, or restricted, did not include non-reef habitat, lacked anthropogenic) is that they serve as nursery areas explicit habitat data, or resulted from indirect for juvenile fishes. The deep reefs of the SEUS sampling methods (Husebø et al., 2002; Uiblein represented a large departure from this pattern as et al., 2003; Auster, 2005; Lorance and Trenkel, we rarely observed or collected small juveniles of 2006; Stone, 2006). Despite compilation from a most fish species. One explanation could be that we variety of video methods not specifically designed missed early recruitment phases because we sampled for fish surveys, the extensive data of Costello et al. in summer or fall; however, it seems unlikely that all (2005) from the northeastern Atlantic provide some species would have the same seasonal patterns of of the best comparisons with our data. They recruitment or that individuals would grow from reported low species richness (5–12 species per site) settlement sizes to the sizes we observed without our across eight sites, with most individuals and species detecting intermediate sizes. Costello et al. (2005) associated with reef or near reef rubble/debris also reported a lack of juvenile fishes from Lophelia zones. Scorpaenids and gadioids were common in reefs, which they attributed to cryptic behavior. It reef habitats, macrourids and morids were common seems unlikely that juveniles of multiple species in transition type habitats, and rajids and Synapho- could completely escape detection. Our frequent branchus spp. were prevalent away from reefs, rotenone stations on the SEUS deep reefs never patterns generally similar to the SEUS. Costello flushed juvenile fishes from hiding. Sulak et al. (in et al. (2005) argued that three-dimensional habitats press) attributed the lack of juvenile fishes on Gulf in the deep-sea of the northeastern Atlantic attract of Mexico deep coral sites to an inability to visually and concentrate fishes, yet no species are restricted feed because of low light levels; however, this to reef habitat. Sulak et al. (in press) also reported a hypothesis lacks evidence. In fact, juveniles of most depauperate fish fauna on a deep Lophelia reef in species are lacking from shelf edge reefs in the the Gulf of Mexico, with some species that seemed SEUS, where ambient light is plentiful (Quattrini restricted to deep reef environments, but they and Ross, 2006). It is not uncommon for juvenile surveyed only a small geographic area. Determining fishes to occupy habitats different from those of the the geographic and bathymetric variability of deep adults (Day et al., 1989; Jones, 1991), and deep-sea reef habitat usage awaits additional data. fishes, like those using SEUS deep reefs, may have The ecological advantages of reef structural life history strategies that place juveniles in habitats complexity (reduction of competition and preda- different from those of adults (Mead et al., 1964). tion, influences on water column physics) that Juveniles of several deep-water species (e.g., operate on shallow reefs (Almany, 2004) should P. americanus, N. bairdii) occupy surface or also exist in deep water. The physical structure of mesopelagic environments (Sedberry et al., 1994, deep coral habitats often rivals that of shallow 1998; Merrett and Barnes, 1996; Haedrich, 1997). It systems, with dense coral matrices (or rocky ledges) is also possible that juvenile fishes inhabit open, soft full of interstitial spaces forming high profile and bottom habitats away from deep reefs, where providing substrata for other sessile invertebrates. predation may be less severe. We collected juveniles The growing reef alters the physics of the water of a few species (Hoplostethus spp., L. barbatulum, column, acceleratingAuthor's bottom currents and enhan- personaland Nezumia spp.) in ottercopy trawls during surveys of cing the environment for corals and attached filter- off reef habitat. feeders (Genin et al., 1986; pers. obs.). The less Most of the deep-sea bottom is composed of soft complex transition reef habitat appears similar (i.e., sediments (increasingly so with depth), a relatively lower profiles, smaller habitat units, more inter- simple, unstructured habitat (Gage and Tyler, spersed soft substrate) to the patch reef and back 1991). Thus, there may be depth and geographic reef formations common to shallow coral reef limits to which deep reef fishes are restricted simply ARTICLE IN PRESS 1002 S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 due to lack of reef habitat. Beyond these limits fish patterns over this SEUS area (Ross and Quattrini, habitat use may be more opportunistic; however, manuscript in preparation), but their usage of deep such a conclusion seems premature, as most of the reef habitat is predictable, not random. Although deep-sea remains poorly explored. Seamounts in data on species interactions (competition, preda- particular are an abundant and under-studied tion) are lacking, we propose that the fishes strongly environment where deep reef fauna may thrive associated with deep coral (or hard bottom) habitats (Rogers, 1994). Aside from the availability of in this study constitute a deep reef community. In structure, the depth limit of deep reef ecosystems contrast to Haedrich and Merrett’s (1990) inter- could be related to the exponential decline in pretation of Mills (1969), our data are consistent transport of labile carbon to the bottom and the with Mills’ definition of community as ‘‘a group of decline of detrital quality with increasing depth organisms occurring in a particular environment, (Carney, 2005), where at some depth there is too presumably interacting with each other and with the little food to support extensive, structure forming, environment, and separable by means of ecological sessile anthozoan or sponge colonies. More explicit survey from other groups.’’ Perhaps without the data on habitats occupied by deep-sea fishes would strong organizing influence of physical structure, help determine whether certain species are merely deep-sea fishes occurring on soft bottoms, especially concentrated by structure, are only present periodi- beyond slope depths, in fact, are just loose cally, or whether they are restricted to these areas. aggregations (Haedrich and Merrett, 1990); how- The concept of fish communities (co-adapted, ever, this hypothesis awaits more detailed data interacting groups) in the deep-sea was challenged, (Koslow, 1993). We hope that this study encourages to be replaced with the proposal that there are only research on deep-sea fish distributions that incor- loose aggregations of individuals whose distribu- porates more explicit data on fish behavior and tions are independent of one another (Haedrich and habitat usage. Merrett, 1990; Haedrich, 1997). A lack of cohesive, identifiable communities could suggest that strong Acknowledgments associations with particular habitats are unlikely. The premise that marine communities may be NOAA Office of Ocean Exploration (Grants nothing more than species collected together, a NA16RP2696, NA030AR4600090, NA040AR4600056, fortuitous or random assemblage, (Haedrich and NA050AR4601065 to S.W. Ross, lead PI) largely Merrett, 1990) was derived from sampling methods supported fieldwork and some data analyses. Uni- that integrated all species caught and obscured ted States Geological Survey (USGS, through the small scale patterns. Koslow (1993) suggested that State Partnership Program) and Minerals Manage- Haedrich and Merrett’s conclusions were based on ment Service (MMS) contributed funds to help with inadequate data and analyses, and upon reanalysis analyses, and we thank Dr. Gary Brewer (USGS) of some of these data species distribution patterns and Greg Boland (MMS) for facilitating the above were revealed. Various collections (usually by support from their agencies. USGS Florida Inte- trawls) have been used to represent, perhaps grated Science Center (through Dr. K.J. Sulak) inappropriately, deep-sea benthic fish distributions. provided a variety of personnel and logistics Obviously, a trawl sample delivers a catch of fishes support for field operations. We thank Dr. M.S. which group together in terms of geography or Nizinski, Dr. J.H. Caruso, Dr. J.V. Gartner, Jr., Dr. depth (assuming the trawl did not cover vertical J.A. Moore, Dr. C.L. Morrison, Dr. D.A. Didier extremes), but whether the species caught formed Dagit, Dr. W.N. Eschmeyer, Dr. T.A. Munroe, Dr. habitat-related patterns would not be discernable. W.D. Anderson, Dr. D.G. Smith, Dr. J.G. Nielsen, We doubt that this inefficient, indiscriminate Dr. B. Fernholm, Dr. J.E. McCosker, and G.H. methodology is adequate for determining the Burgess for helpful discussions, assistance with existence of deep-seaAuthor's fish communities. We hy-personalspecies identifications, copy and/or participation in pothesize that some slope fishes have experienced cruises. Dr. D.M. Wyanski provided information similar selective pressures controlling habitat usage on the South Carolina E. minor specimen. M.S. and distributions as shallow species and responded Nizinski and T.A. Munroe provided helpful reviews to them in ways leading to communities closely tied of this paper. E. Baird, A. Howard, T.L. Casazza, to structured habitat. The deep reef fishes we A.M. Necaise, and J.P. McClain assisted with identified exhibited some variation in distributional various aspects of this project, and B. York helped ARTICLE IN PRESS S.W. Ross, A.M. Quattrini / Deep-Sea Research I 54 (2007) 975–1007 1003 with data editing and organization. M.L. Partyka Burgess, G.H., Link, G.W., Ross, S.W., 1979. Additional marine assisted with preparation of maps and cruise fishes new or rare to Carolina waters. Northeast Gulf Science activities. We thank Harbor Branch Oceanographic 3, 74–87. Cailliet, G.M., Andrews, A.H., Wakefield, W., Moreno, G., Inst. ship, submersible and shore-based personnel Rohdes, K.L., 1999. Fish faunal and habitat analyses using for assisting with numerous cruises. The 2001 and trawls, camera sleds and submersibles in benthic deep-sea 2006 R.V. Cape Hatteras missions were sponsored habitats off central California. Oceanologica Acta 22, by the Duke/UNC Oceanographic Consortium (to 579–592. S.W. Ross), and we thank that ship’s personnel for Carney, R.S., 2005. 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Author's personal copy
OTC 18510
AUV-Based Environmental Characterization of Deep-Water Coral Mounds in the Straits of Florida M. Grasmueck, G.P. Eberli, T.B.S. Correa, D.A. Viggiano, J.Luo, RSMAS University of Miami; G.J. Wyatt, Quester Tangent; J.K.Reed, A.E. Wright, and S.A. Pomponi, Harbor Branch Oceanographic Institution.
Copyright 2007, Offshore Technology Conference instrumentation has focused most research activity and related This paper was prepared for presentation at the 2007 Offshore Technology Conference held in discoveries of deep-water coral habitats to the north and Houston, Texas, U.S.A., 30 April–3 May 2007. central Atlantic, the Gulf of Mexico and the north-east This paper was selected for presentation by an OTC Program Committee following review of 4,5 information contained in an abstract submitted by the author(s). Contents of the paper, as Pacific . In the Straits of Florida, abundant mound-forming presented, have not been reviewed by the Offshore Technology Conference and are subject to corals in water depths of 400–800 m have been documented in correction by the author(s). The material, as presented, does not necessarily reflect any position of the Offshore Technology Conference, its officers, or members. Papers presented at over 40 years of dredge sampling, submersible dives and OTC are subject to publication review by Sponsor Society Committees of the Offshore 6-12 Technology Conference. Electronic reproduction, distribution, or storage of any part of this seismic acquisition (Figure 1). This extensive collection of paper for commercial purposes without the written consent of the Offshore Technology samples and observations however can not be put into a Conference is prohibited. Permission to reproduce in print is restricted to an abstract of not more than 300 words; illustrations may not be copied. The abstract must contain conspicuous geomorphologic context as existing bathymetric charts do not acknowledgment of where and by whom the paper was presented. Write Librarian, OTC, P.O. Box 833836, Richardson, TX 75083-3836, U.S.A., fax 01-972-952-9435. resolve coral mounds. Such sparse information has proven inadequate to answer questions in regards to mound Abstract morphology, bottom current dynamics and nutrient sources The first AUV survey across five fields of deep-water coral supporting life at these depths. Furthermore the limited data mounds in the Straits of Florida reveals an unexpected high set has so far prevented assessment of the biodiversity and the abundance and variability of mounds in water depths of 590 – potential need for protection from over-fishing and underwater 875 m. A drop camera and a series of dives with the Johnson- construction. High-resolution maps of morphology and Sea-Link submersible confirmed living corals on each of the oceanographic conditions resolving features at the 1-10 m five investigated sites. The morphology of the mounds is scale are a basic requirement to make further progress in highly diverse, ranging from isolated mounds to well- understanding deep-water coral mound distribution and developed ridges with more than 100 m of relief. Along the genesis. toe-of-slope of western Great Bahama Bank antecedent Autonomous Underwater Vehicles (AUV) bring an topography seems to be the controlling factor for mound integrated suite of mapping and oceanographic sensors close location while further west currents appear to control the to the seabed for high-resolution data acquisition and give the formation of ridges. The comprehensive suite of sensors on opportunity to fill the scale gap of basic information in deep- board the AUV allows correlation of geophysical parameters water environments. Described here are the initial results of a and oceanographic observations. Acoustic Doppler current 7 day cruise during which the AUV mapped five deep-water coral mound fields in the Straits of Florida covering a total meter data document three different bottom current regimes 2 consisting of unidirectional or bi-directional tidal flow. The bi- area of 130 km . Acquiring the same amount of data without directional current pattern is not visible on backscatter data an AUV would have taken multiple cruises and possibly and only vaguely reflected in the mound morphology. In areas several years with costs exceeding the single AUV mapping of uniform current direction mounds face the currents and cruise. By simultaneously acquiring a comprehensive suite of align perpendicular to the current to form long ridges and high-resolution seabed and oceanographic data the dynamic intervening troughs. The synoptic seabed and oceanographic and complex environment spanning entire coral mound fields data recorded by the AUV characterize the dynamic and can be assessed for the first time. We selected five mapping complex environments of entire coral mound fields at a sites off the Miami Terrace, in the center of the Straits of resolution of 1–3 m. Florida and at the toe-of-slope of Great Bahama Bank. Mound distribution, morphology and currents are different for each Introduction survey site. Subsequent groundtruthing with a drop camera Deep and cold-water coral ecosystems are less known but and a submersible found corals in all five areas. By comparing more widespread than their warm-water counterparts restricted the submersible observations with the AUV data the to shallow tropical seas1,2. Cold-water corals and associated distribution of deep-water coral habitats in each area can be fauna flourish in oceanic waters of all latitudes at depths of predicted. This comprehensive dataset allows us to assess the several hundred to over one thousand meters with relationship between mound morphology and current direction temperatures between 4° and 12° C and require no sunlight3. and the abundance and distribution of corals on the mounds. The limited availability and high cost of deep-water 2 M. Grasmueck et al. OTC 18510
Figure 1. Seafloor morphology of the Straits of Florida based on National Ocean Service Hydrographic Survey Data. This major seaway connects the Gulf of Mexico with the Atlantic. It is bordered by the Florida peninsula in the NW, Cuba in the south, and the Bahamas in the east. The Florida Current, a warm surface current, flows through the Straits of Florida into the North Atlantic where it converges with the smaller Antilles Current to form the Gulf Stream. The Florida Current has produced stream line shaped drift deposits. Superimposed on the map are the locations of scientific dredge sampling in the 1960s and 70s at water depths deeper than 400 m 8,11,19. The dredge locations highlighted with red dots retrieved mound-forming deep-water corals Lophelia pertusa, Enallopsammia profunda and Madrepora oculata. The five areas mapped with the AUV are marked with black and white rectangles
Survey Methods
Sea Surface-Based Multibeam Reconnaissance Mapping. To make the best use of the AUV for mapping coral mound fields and avoiding featureless flat seabed areas we performed sea surface-based multibeam bathymetry surveys just before launching the AUV. The hull-mounted 12 KHz EM120 multibeam system on board the R/V NORTHERN RESOLUTION (Figure 2) produced 50 m gridded bathymetric maps resolving mound structures not visible on the best available bathymetric chart of the Straits of Florida (Figure 1). As the dredge location coordinates have large position uncertainties of where exactly the samples were retrieved from the seabed, the reconnaissance multibeam mapping was essential to locate the coral mound fields. The dredge coordinate locations served as a starting point for Figure2. The R/V NORTHERN RESOLUTION used for hull- scanning the seabed at 10 kts vessel speed. The multibeam mounted multibeam reconnaissance mapping and AUV swathwith was 3.5 km at 800 m water depth. The resulting deployment preliminary maps (Figure 3) served as the basis for designing and planning the tracks of the subsequent AUV surveys.
OTC 18510 AUV-Based Environmental Characterization 3
Positioning and navigation accuracy of the AUV at 800 m depth is better than 3 m 13. A Kalman filter combines the inputs from the AUV fiber optic gyro compass, acoustic Doppler profiler and high precision pressure (depth) sensor. Positioning drift is minimized with fixes from an ultra short baseline (USBL) acoustic and differential global positioning system installed on the mother vessel14,15. Communications between mother vessel and AUV by acoustic modem allow quality control on decimated data in real time during the mission. The high-resolution bathymetry including backscatter amplitudes were acquired using a 200 kHz EM2000 multibeam system installed on the AUV. The swath width is 300 m providing 100 m overlap between adjacent lines. The multibeam data were corrected for heave, pitch and roll monitored by precision accelerometers. The data were processed on board the R/V NORTHERN RESOLUTION into bathymetric maps with 3 m resolution and acoustic backscatter maps with 1 m resolution within a couple hours after the AUV had resurfaced. Average bottom current velocity and direction in the 40 m water column between AUV and seafloor were extracted from the acoustic doppler profiler installed on the AUV. Salinity and water temperature were recorded at 1 s intervals for later fusion with the position data and plotting along the AUV track lines, monitoring how oceanographic conditions evolved during the survey.
Figure 3. EM120 multibeam bathymetry acquired for Figure 4. The AUV resting on the retractable launch and retrieval reconnaissance of Site 1. The white stippled rectangle shows the area selected for high-resolution AUV mapping. The 50 m system at the stern of the R/V NORTHERN RESOLUTION. resolution map images numerous mound features associated with low relief ridges extending in a slightly divergent pattern in Groundtruthing with Drop Camera and JSL Submersible. E-W direction The 40 m cruising altitude of the AUV is too high to illuminate the seabed for still pictures or video to verify the High-Resolution Data Acquisition with the AUV cruising presence of corals on the mounds. Therefore, we developed a 40 m above the Seabed. low-cost drop camera to get a first glimpse of corals The Autonomous Underwater Vehicle (AUV) was immediately after the AUV mapping was completed. The drop deployed from the stern of R/V NORTHERN RESOLUTION. camera consisted of a compact digital camera within a 15 cm Over 7 days in December 2005, the Hugin 3000 AUV (Figure diameter evacuated glass sphere rated to 6700 m depth taking 2 4) mapped five sites ranging from 14–48 km in 590–875 m pictures every 1.5 seconds for 70 minutes (Figure 5). The water acquiring 1–3 m resolution multibeam data together camera is kept neutrally buoyant 1–2 meters above the with sidescan data, subbottom profiles, current vectors, seafloor by the variable weight of a loose steel chain. LED salinity, temperature and methane content. The AUV is lights illuminate the seafloor. An acoustic pinger attached 50 powered by an aluminum oxygen fuel cell providing a mission m above the camera enabled ± 5m accurate USBL tracking. endurance of up to 55 hours and cruises 40 m above the With the 50 m of line between pinger and camera the seabed seafloor at a speed of 1.8 m/s scanning the seafloor in parallel position of the photographs taken is known to within about 50 lines spaced 200 m in similar fashion to a lawn mower. m radius. The AUV bathymetric map was used to select an 4 M. Grasmueck et al. OTC 18510 off-mound camera landing site and to steer the camera trajectory during the deployment. Currents and the vessel pulling on the line moved the camera at an approximate speed of 0.5 m/s along the seabed. We only deployed the drop camera once during the AUV cruise in order to save valuable AUV time. The drop camera deployment was, however, a proof of concept for future inexpensive groundtruth missions from small boats before or after AUV surveys.
Figure 6. The JOHNSON-SEA-LINK (JSL) submersible carries four people: Two in the acrylic capsule up front and two in an aluminum pressure chamber in the back.
Results
Site 1: Mounds aligned along off-Bank Ridges at the Toe- of-Slope of Great Bahama Bank. This site in 590-710 m water depth is just north of the ODP Site 1007 drilling location at the base-of-slope along the western margin of Great Bahama Bank. Several mound features protruding from the seafloor had been noticed on
Figure 5. Compact drop camera developed for groundtruthing of some of the high-resolution 2D seismic lines acquired during coral mounds from small boats. the site survey for Ocean Drilling Program (ODP) Leg 16617. Although the seismic data detected mounds, the 600 m line Harbor Branch Oceanographic Institution’s (HBOI) spacing did not provide information about the abundance and JOHNSON-SEA-LINK (JSL) submersible dives launched distribution of the mounds. The new 3 m grid resolution AUV from R/V SEWARD JOHNSON some months after the AUV bathymetry map now shows 37 mounds, which are at least 25 cruise provided us with a more extensive and precise m high (Figure 7). The tallest mound reaches a height of 85 m groundtruthing of mound coverage at all 5 AUV survey sites and has a minimum base width of 350 m. The drop camera (Figure 6). The JSL submersible carries four persons to a deployment across this mound revealed living corals near the maximum depth of 1000 m. It has an acrylic sphere that peak and coral rubble with sponges and brittle stars on the provides > 180º visibility to the pilot and the observer in the north facing slope. The AUV map also images more than 180 front. Color videotapes were recorded along the dive tracks smaller mounds. The distribution of mounds is non-random with an external pan and tilt video camera. Submersible and follows low relief ridges (500-1500 m wide and 5 m high) navigation used an USBL positioning system and calculated extending in a slightly divergent pattern in an E-W direction the submersible’s real-time position throughout each dive. (Figure 3). The shapes of the mounds are diverse, including Analysis of USBL tracking accuracy for a worst-case tracking single conical, pyramid or wedge shaped peaks, twin peaks, scenario estimated a maximum statistical positioning error of and an almost perfect heart shape. Fields of 2-4 m tall ridges 9.6 m at a depth of 500 m (J. Kloske, Florida Institute of spaced by 15-25 m occur at the base of most large mounds. 16 Oceanography, pers. comm.; and ). A Geographical Some mounds are completely or partly surrounded by 5-10 m Information System (GIS) was used to overlay the deep and 25-50 m wide moats. Smaller mounds generally lack submersible track positions and time stamps onto the AUV the moat but have a sediment wedge attached that thins acquired bathymetry, backscatter and sidescan sonar maps. towards the north. The timestamps on the video frames allowed a direct The bottom currents within the 40 m water column correlation of visual observations and related patterns on the between AUV and seafloor showed a remarkable variation in AUV maps. strength and direction18. The currents changed direction between north- and southward flow seven times over 45 hours. The average time interval between changes was approximately 6 hrs. Current strength never exceeded 0.5 m/s with sustained peak strengths of 0.2 m/s in north or south direction. The change of direction every six hours indicates tidal control on the currents at the toe-of-slope of Great Bahama Bank in 590- OTC 18510 AUV-Based Environmental Characterization 5
710 m water depth. This current behavior is rather surprising direction appears to be slightly prevailing. This is in as the surface currents flow northward. The bottom currents agreement with the consistent northward orientation of the obviously are decoupled from the surface currents and sediment wedges associated with small mounds. dominated by diurnal tides. The north-flowing current
Figure 7. Site 1: High-resolution bathymetry based on 200 kHz multibeam data acquired with the AUV at a cruising altitude of 40 m above the seafloor. The map images more than 200 mounds within an area of 48 km2 at the toe-of-slope of Great Bahama Bank. The shapes of the mounds are diverse, including single conical, pyramid or wedge shaped peaks, twin peaks, random shaped mounds and even an almost perfect heart shape. Insets are zoomed up views of the largest mounds with a height of 85 m and photographs of coral cover taken with the drop camera.
Site 2: Mounds related to Slump Features at Toe-of-Slope small-scale ridges lining the tip and northern edge of the of Great Bahama Bank. elongate plateau. The backscatter data show stronger seabed Following the toe-of-slope 40 km to the north from Site 1 reflection amplitudes from these ridges than the surrounding we encountered a section with a steep escarpment reaching seabed. Based on the submersible dive, the higher reflectivity heights of over 100 m (Figure 8). The escarpment follows two is interpreted to be caused by living- and/or dead coral on large scars and a 3.2 km long and up to 0.9 km wide plateau these ridges. The acoustically softer seabed between the ridges that prominently raises 40-150 m from the surrounding gently and in the southern portion of the plateau are likely composed sloping seabed. The JSL submersible dive I-4812 visiting this of muddy sediments. Numerous mounds features are also elongate plateau in 2005 encountered 1.0–1.2 m high living visible to the south-west of the plateau. Backscatter Lophelia coral thickets near the peaks of NE-SW oriented amplitudes suggest coral cover and soft sediment surrounding ridges in 690 m of water. The slopes of the coral ridges were the mounds. For some mounds the shapes of the darker covered with 100% coral rubble and some 1.0 m dead standing colored backscatter anomalies do not follow the contours of coral thickets. The AUV multibeam data show numerous the mounds but extend over the flat seabed around and 6 M. Grasmueck et al. OTC 18510 between the mounds, indicating biological growth and rubble into the current feature darker backscatter amplitudes than the production. These mounds have not yet been groundtruthed. It down current side and adjacent troughs. The JSL submersible appears that the elongate plateau is a remnant between two dives confirmed bio-accumulation in form of coral rubble and slump scars originating from basal slope instability of Great standing dead and living corals and associated fauna on the Bahama Bank. The foundations for deep-water corals on the slopes facing the current and muddy sediments in the lee-side group of mounds in front of the escarpment are likely the troughs. blocks from this mass gravity flow. The bottom current data from the AUV survey show a north-south reversal indicating a Site 5: Sinusoidal Ridges at the Base of the Miami Terrace. bi-directional tidal current regime as already encountered at Neumann and Ball (1970) reported from their 1969 the first site with maximum current strengths reaching 0.25 Aluminaut submersible dive “….at the base of the Miami m/s. Terrace, is an elongate trough. The bottom here is characterized by ridges and mounds of muddy sand capped by Site 3: Coalesced Mounds form off-Bank Ridges at the thickets of living deep-water branching coral.” The new AUV Toe-of-Slope of Great Bahama Bank. bathymetric map (3.8 x 7.1 km) shows how coral mounds at Site 3 is also on the base-of-slope of Great Bahama Bank the toe-of-slope of the Miami Terrace (875 to 660 m) are in 710 to 840 m water depth. The site is located 15 km west of developed as a series of sinusoidal, asymmetrical ridges that Bimini and 80 km north from the Site 2. In 1964 R/V GERDA are approximately 800 m long, less than 30 m high, and 100 to retrieved Enallopsammia profunda corals in one dredge 400 m wide (Figure 12). The AUV bottom current data sample (Sample g29719). The high-resolution bathymetric map confirm consistent southward bottom flow (opposite to the covering 3.8 x 3.6 km area shows 4 major ridges of coalesced surface current) as Neuman and Ball had already experienced mounds aligned in NE-SW direction (Figure 9). The largest during their dive. The ridges are steeper with stronger ridge is 0.5 km wide and 2 km long. The tallest peak on the (=darker) backscatter on their northern sides that face the ridge is 120 m high and has a pyramidal shape. The 2006 JSL currents. The 2006 JSL submersible dive encountered deep- submersible dive on this mound found dense living coral water coral rubble and thickets on the north facing slopes and thickets on the flank and at the peak. In addition to the crests alternating with a muddy seabed in the leeward troughs. coalesced mound ridges, more than 150 individual mounds are At the base of the slope the coral coverage ends abruptly as it distributed throughout the site. Some of the mounds have transitions sharply into a barren sand dune field devoid of pronounced elliptic scour depressions on the north side while corals. In this sand dune field, the steeper slopes of the sand others have elongate sediment wedges attached. The acoustic waves face to the south as expected from loose sand backscatter map shows linear dark striations originating at the transported by south flowing bottom currents. mounds with some extending to over 1 km in a NNW direction (Figure 10). These striations are nearly Discussion perpendicular to the orientation of the mound ridges. The currents measured during the AUV survey indicate tidal AUV provides Synoptic View of entire Coral Mound reversals every 6 hours in a north or south direction with the Fields. north flowing currents slightly prevailing and a sustained Previous studies of the deep-water coral mounds in the strength of 0.3 m/s. Straits of Florida relied primarily on data acquired from a single mound perspective with a limited number of sensors. Site 4: Chevron Pattern Mound Clusters in the Center of The 130 km2 of bathymetric data acquired by the AUV Straits. provides high-resolution maps of entire mound fields. This site is located 30 km offshore from Bimini in the Furthermore, together with the multibeam backscatter and center of the Straits of Florida. Three R/V GERDA dredges side-scan swaths bottom type and coverage can be assessed (Samples g311, g317, g35419) from this area retrieved field wide. There is a close association between high Lophelia pertusa and Enallopsammia profunda. The AUV backscatter patches (dark shades on backscatter plots) with mapped an area of approximately 3.2 x 4.2 km. Nearly 70% of biologic accumulations composed of coral rubble, standing the area is covered with small mounds, most less than 5 m dead and live corals. On the basis of this association verified high, producing a knobby seafloor topography (Figure 11). In at 5 sites we can now predict with a high degree of certainty the center of the study area the mounds are coalesced to form where such deep water coral habitats occur on multibeam hills and ridges of approximately 40 m height. Individual maps. The new 1-3 m grid resolution maps now allow precise mounds have a horseshoe shape with the convex side facing planning of submersible or ROV tracks for efficient towards the S and SSE. The backscatter map shows lineations groundtruthing at critical locations. The resolution of the attached to the mounds which trend in a NNW direction. The seabed maps has proven adequate to georeference bottom currents recorded over the 13 hour AUV survey time observations made within the illuminated area of the JSL were consistently flowing to the north with an average submersible. velocity of 0.23 m/s (maximum 0.43 m/s) with azimuths varying between 330° and 50°. Many knobs are aligned and Surprising Diversity and Abundance of Deep-Water form chains oriented at an angle of 70° – 90° with the current Corals in the Straits of Florida. direction. The knob chains form a characteristic chevron Mound morphology is not uniform, even for mounds in pattern very different from the ridge morphology observed at close proximity to each other. Mound morphologies range Site 3 just 15 km to the east. The slopes of the mounds facing from predominantly isolated structures (Site 1) to enormous, OTC 18510 AUV-Based Environmental Characterization 7 well-developed ridges with more than 100 m of relief (Site 3). 5) The majority of chains and ridges of coral mounds are At these two sites on the toe-of-slope of western Great aligned perpendicular to the prevailing current. Bahama Bank the mounds are aligned with low-relief These observations partly corroborate but also contradict sediment ridges and lows, characteristic of the lower slope earlier interpretations. For example, the fact that mounds face morphology surrounding carbonate platforms20. The mound into the current has already been noted in early submersible distribution seems to follow the sediment ridges that trend observations21. We found no good examples of the “tear drop” approximately perpendicular to the platform margin. This morphology with the long mound axis oriented parallel to the configuration is also perpendicular to the tidal-controlled prevailing current direction10. Our data show no correlation current that flows either north or south. Similarly mound between mound height and current strength. The tallest morphology seems not to be controlled by the current mounds are situated in areas where the current is tidally direction as mounds display a wide variability of shapes. dominated and switches directions every six hours. The style of coral mound architecture changes dramatically across the Straits of Florida from Bimini to Miami. The Survey Strategy for efficient Characterization of the entire knobby mounds in the middle of the Straits of Florida at Site 4 Florida Straits and other Deep-water Habitat Regions. are densely clustered and form a chevron pattern. At the base The new AUV surveys cover only a minuscule portion of of the Miami Terrace the mounds form sinusoidal ridges the Straits of Florida. Time and cost would be prohibitive for trending perpendicular to the currents. The boundaries mapping the entire Straits with an AUV. As an outcome from between coral mound fields and featureless muddy or sandy this first application of AUV technology to mapping of deep- seafloor are sharp. water coral environments we propose the following survey Deep-water corals and related fauna exhibit a remarkable strategy, which is also applicable to deep-water habitats adaptability to different environmental conditions by changes elsewhere: 1) Regional surface bathymetric mapping and in mound morphology and configuration. It also appears that backscatter classification for identification of mound fields. 2) previous estimates of number of mounds and extent of deep- Drop camera deployment from small boats on selected water coral habitats will have to increase by orders of mounds to determine presence of live or dead coral cover. 3) magnitude. The five areas mapped with the AUV contain High-resolution AUV surveying of key mound fields. 4) hundreds of coral mounds with heights of 1–120 m covering Based on the AUV maps perform further drop camera 30% of the 130 km2 total area mapped. The abundance of deployments for more detailed groundtruth. 5) Use of deep-water corals was confirmed by subsequent visual submersibles and ROV for sample collection and deployment groundtruthing with the drop camera and the human-occupied of permanent monitoring equipment. This scaled approach submersible. The previously underestimated abundance and including modern oceanographic mapping tools requires a new diversity of modern deep-water coral mounds is likely to hold level of interdisciplinary deep-water research, unifying local true for the geologic record as well. As the AUV mapping observations with their regional environments. Such areas were specifically selected for potential coral coverage collaboration together with an extensive mapping strategy will the mound and area coverage numbers derived from this study be needed to resolve questions related to nutrient supplies and do not apply to the entire Straits of Florida. the seemingly high mortality of the deep-water coral mounds in the Straits of Florida. New Insights on Influence of Currents on Coral Mound Morphology and Distribution. Conclusion Mound distribution, morphology and currents are different In just a few days the AUV acquired the high-resolution for each of the five mapped sites. Taken together the AUV morphologic and oceanographic maps essential for the data reveal some general trends: comprehensive environmental assessment of five different 1) Bottom currents are not unidirectional in the Straits of deep-water coral mound fields. The AUV data provide for the Florida. Tidal dominated currents along the western slope of first time a field-wide perspective not obtainable by dredge Great Bahama Bank flow both north and southward. In the sampling or visual observation from remotely operated center of the Straits flow is northward while along the slope of vehicles or submersibles. Each of the five sites revealed a the Miami Terrace the flow is southward. Sediment wedges distinct character in terms of mound morphology, that are attached to mounds as well as the striation patterns configuration and bottom currents. Most surprising are the visible on backscatter maps do not show the tidal reversing as high abundance of coral mounds at all sites and the variability they are pointing in one direction only. Furthermore, the AUV of both the distribution and the shape of individual mounds. current data display directional variability, which is also not Many coral mounds align or coalesce along linear ridges seen in the backscatter striation patterns. approximately perpendicular to the prevailing current 2) Coral mounds grow taller in bi-directional tidal bottom direction. Tear drop shape of individual mounds is the current regimes. exception rather than the rule. Similar variability and local 3) Chains of closely spaced smaller coral mounds develop abundance of deep-water corals is expected at other locations. as a response to uniform bottom current direction. AUVs are efficient high-resolution mapping platforms which 4) Corals grow into the currents creating steeper mound can guide future sampling and monitoring to critical locations slopes facing the current, indicating that coral growth and not and provide the base maps for making inventories and sediment accumulation from traction current control the shape management plans of deep-water habitats. of the mounds.
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Acknowledgments hardgrounds in the northeastern Straits of Florida, Palaios, 5, We received ship time for the AUV survey from the NOAA 15–33. Ocean Explorer program. Justin Manley (OE program 11. Cairns, S., (2000), A revision of the shallow-water manager), Heather Langill (C&C Geophysical Operations & azooxanthellate scleractinia of the Western Atlantic, Studies of the natural history of the Caribbean Region, 75, 240. Bid Coordinator) and Chas Honea (C&C AUV Field Project 12. Reed, J.K., Weaver, D., and Pomponi, S.A., (2006), Habitat and Manager) were instrumental for facilitating this project. We fauna of deep-water Lophelia pertusa coral reefs off the express our gratitude to the officers and crew of the R/V Southeastern USA: Blake Plateau, Straits of Florida, and Gulf of NORTHERN RESOLUTION and the scientific and technical Mexico, Bulletin of Marine Science, 78, 343–375. staff of C&C Technologies. Their efficiency and competence 13. Jalving, B., Gade, K., Hagen, O.K., and Vestgård, K., (2003), A enabled data acquisition and processing in a very short time. Toolbox of Aiding Techniques for the HUGIN AUV Integrated We especially appreciated how they accommodated all our Inertial Navigation System, Proceedings from Oceans 2003, 8. special requests in their workflow. 14. Chance, T.S., and Northcutt, J.G., (2001), Deep water AUV The crews of the R/V SEWARD JOHNSON, and the experiences, Proceedings of the U.S. Hydrographic Conferences, 8. JOHNSON-SEA-LINK submersibles are gratefully thanked 15. George, R.A. Advances in AUV remote-sensing technology for for their logistical support. A grant provided by the State of imaging deepwater geohazards The Leading Edge, Volume 25, Florida, Medicines from Florida's Oceans Project (HBOI Issue 12, pp. 1478-1483 doi:10.1190/1.2405333 Project Nos. S2156 and S2168) provided funding for these 16. Opderbecke, J. 1997. At-sea calibration of a USBL underwater cruises. We are especially grateful to all the personnel of the vehicle positioning system, Oceans 1997, MTS/IEEE Conf. Division of Biomedical Marine Research at HBOI who are Proc. 1: 721–726. responsible for various aspects of research during these 17. Anselmetti, F.S., Eberli, G.P., and Ding, Z.-D., (2000), From the cruises. Great Bahama Bank into the Straits of Florida: A margin Pre- and post-cruise research was made possible by the architecture controlled by sea level fluctuations and ocean currents, GSA Bulletin, 112, 829–844. Industrial Associates of the Comparative Sedimentology 18. Grasmueck, M. Eberli, G., Viggiano, D.A. Correa, T. Rathwell, Laboratory. Angela D. Rosenberg helped with analyzing the G. and Luo, J., 2006, Autonomous Underwater Vehicle (AUV) submersible videos. IVS 3D Fledermaus™ software was used mapping reveals coral mound distribution, morphology and for rendering and visualization of the bathymetric maps. oceanography in deep water of the Straits of Florida, Geophys. Res. Lett., 33, L23616, 6p, doi:10.1029/2006GL027734. References 19. Cairns, S., 1976, Review of the deep-water ahermatypic corals 1. Paull, C.K., Neumann, A.C., Ende, B.A., Ussler, W., and (Scleractinia) of the tropical western Atlantic, Dissertation Rodriguez, N.M., (2000), Lithoherms on the Florida-Hatteras University of Miami.316p. slope, Marine Geology, 136, 83-101. 20. Mullins, H. T., Heath, K.C., Van Buren, H.M., and Newton, 2. Freiwald, A., Fosså, J.H., Grehan, A. Koslow, T. and Roberts, C.R., (1984), Anatomy of a modern open-ocean carbonate J.M., (2004), Cold-water Coral Reefs, 84 pp., UNEP-WCMC, slope: northern Little Bahama Bank: Sedimentology, 31, 141- Cambridge, UK. 168. 3. Roberts, J.M., Wheeler, A.J., and Freiwald, A., (2006), Reefs of 21. Neumann, A.C., Kofoed, J.W., and Keller, G.H., (1977), the Deep: The Biology and Geology of Cold-Water Coral Lithoherms in the Straits of Florida, Geology, 5(1), 4–10. Ecosystems, Science 28(312), 543–547, doi:10.1126/science.1119861. 4. De Mol, B., Van Rensbergen, P., Pillen, S., Van Herreweghe, K., Van Rooij, D., McDonnell, A., Huvenne, V., Ivanov, M., Swennen, R., and Henriet, J. P., (2002), Large deep-water coral banks in the Porcupine Basin, southwest of Ireland, Marine Geology, 188, 193–231. 5. Ferdelman, T.G., Kano, A., Williams, T., and the IODP
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