Habitat and Fauna of Deep-Water Lophelia Pertusa Coral Reefs Off the Southeastern U.S.: Blake Plateau, Straits of Florida, and Gulf of Mexico

Home , Apo Island, Apo Reef, Chlorophthalmus, French Reef, Hexactinellid, Pronotogrammus

BULLETIN OF MARINE SCIENCE, 78(2): 343–375, 2006 CORAL REEF PAPER

Habitat and Fauna of Deep-Water Lophelia pertusa Coral Reefs off the Southeastern U.S.: Blake Plateau, Straits of Florida, and Gulf of Mexico

John K. Reed, Doug C. Weaver, and Shirley A. Pomponi

Abstract Expeditions from 1999 to 2004 for biomedical research explored various deep-sea coral ecosystems (DSCE) off the southeastern U.S. (Blake Plateau, Straits of Florida, and eastern Gulf of Mexico). Habitat and benthos were documented from 57 dives with human occupied submersibles and three with a remotely operated vehicle (ROV), and resulted in ~100 hrs of videotapes, 259 in situ digital images, 621 museum specimens, and > 400 microbial isolates. These were the first dives to document the habitat, benthic fauna, and fish diversity of some of these poorly known deep- water reefs. Fifty-eight fish species and 142 benthic invertebrate taxa were identified. High-definition topographic SEABEAM maps and echosounder profiles were also produced. Sites included in this report range from South Carolina on the Blake Plateau to the southwestern Florida slope: 1) Stetson Lophelia reefs along the eastern Blake Plateau off South Carolina; 2) Savannah Lophelia lithoherms along the western Blake Plateau off Georgia; 3) east Florida Lophelia reefs, 4) Miami Terrace escarpment in the Straits of Florida; 5) Pourtalès Terrace off the Florida Keys; and 6) west Florida Lophelia lithoherms off the southwestern Florida shelf in the Gulf of Mexico. These are contrasted with the azooxanthellate deep-water Oculina reefs at the shelf-edge off central eastern Florida. The fisheries and biopharmaceutical re- source potential of these deep-water habitats remain relatively unknown. Although these habitats are not currently designated as marine protected areas (MPAs) or coral habitat areas of particular concern (HAPCs), they are ecologically diverse, vulnerable to physical destruction, and irreplaceable resources. Activities involving bottom trawling, pipelines, or oil/gas production could negatively impact these reefs. National Oceanic and Atmospheric Administration (NOAA) Fisheries and the South Atlantic Fishery Management Council are currently developing priority mapping sites of the DSCEs within this region, and these data may provide potential targets for new MPAs and HAPCs.

Deep-sea coral ecosystems (DSCEs) are common off the southeastern U.S. within the Exclusive Economic Zone (EEZ). These include a variety of high-relief, hard-bottom habitats at numerous sites from the Blake Plateau off North Carolina, southward through the Straits of Florida, and in the eastern Gulf of Mexico. However, only a few have been mapped or have had their benthic and fish resources characterized.D eep- water reefs are sometimes defined as coral banks, coral mounds, bioherms, or lithoherms (Teichert, 1958; Stetson et al., 1962; Neumann et al., 1977; Wilson, 1979; Reed, 1980, 2002a,b; Freiwald et al., 1997; Paull et al., 2000; Fosså et al., 2002; Reed et al., 2005a,b). In general, deep-water banks occur below the effects of waves and the corals lack symbiotic algae (zooxanthellae). A bioherm is a deep-water coral bank that over centuries has formed a mound of unconsolidated sediment and coral debris and is capped with thickets of coral, such as Oculina or Lophelia (Reed, 2002a,b), whereas lithoherms are high-relief, lithified carbonate mounds, rather than unconsolidated sediment mounds, and also may be covered with thickets of live coral (Neumann et al.,

1977). Rogers (1999) suggested that deep-water coral banks fall within the definition of a coral reef based on their physical and biological characteristics. The dominant corals on deep-water reefs in this region are the azooxanthellate, colonial scleractinian hard corals, Oculina varicosa, Lophelia pertusa , Enallopsam- mia profunda, Madrepora oculata, and Solenosmilia variabilis (Reed, 2002a,b). Numerous solitary coral species are also common (Cairns, 1979, 2000) along with numerous species of calcified hydrozoans (family Stylasteridae), bamboo octocorals (family Isididae), and black corals (order Antipatharia). These reefs provide hard- bottom substrate and habitat for sessile macrofauna including scleractinian corals, octocorals (gorgonians), black corals, and sponges, which in turn provide habitat and living space for a relatively unknown but biologically rich and diverse community of associated fishes, crustaceans, mollusks, echinoderms, polychaete and sipunculan worms, and other macrofauna. Lophelia, Enallopsammia, and Madrepora form reefs in 490–900 m on the Blake Plateau from North Carolina to Florida and in the Straits of Florida (Stetson et al., 1962; Milliman et al., 1967; Uchupi, 1968, 1969; Neumann and Ball, 1970; Emery and Uchupi, 1972; Menzies et al., 1973; Mullins et al., 1981; Messing et al., 1990; Sedberry, 2001; Reed, 2002b; Reed and Ross, 2005). In addition, azooxanthellate, deep-water Oculina coral reefs occur in 70–100 m along the shelf-edge off central eastern Florida (Avent et al., 1977; Reed, 1980, 2002a; Reed et al., 2005b). The high-relief limestone outcrops, escarpments, and bioherms of the Miami and Pourtalès Terraces off southeastern Florida are not Lophelia reefs, per se, but do provide hard-bottom habitat for corals, octocorals, and sponges (Neumann and Ball, 1970; Ballard and Uchupi, 1971; Gomberg, 1976; Reed et al., 2005a). Lophelia bioherms are also present at the base of the Miami Terrace escarpment (~670 m) within the axis of the Straits of Florida, but little is known of their distribution, abundance, or associated fauna (Neumann and Ball, 1970; Cairns, 1979). In the Gulf of Mexico on the southwest Florida slope, a 20-km long zone of high-relief (10–15 m) Lophelia lithoherms at 500 m provides habitat for coral and hard-bottom communities (Newton et al., 1987; Reed et al., 2004). In addition, fairly extensive Lophelia thickets grow on Viosca Knoll in the northern Gulf of Mexico off Alabama (Schroeder, 2002; Reed et al., 2004), and occur in DeSoto Canyon off Ala- bama and Green Canyon off Louisiana (Schroeder et al., 2005). As a result of recent research expeditions (1999–2004) by Harbor Branch Oceano- graphic Institution (HBOI) for biomedical research using human occupied submersibles and remotely operated vehicles (ROV), information has been compiled on the status, distribution, habitat, and biodiversity of some of these relatively unknown and newly discovered deep reef ecosystems. We located and mapped nearly 300 mounds during echosounder transects and Johnson-Sea-Link (JSL) submersible dives off the east coast of Florida, many of which proved to be newly discovered deep-water reefs, up to 168 m in height and at depths of 700–800 m. Submersibles and ROVs were used to ground-truth additional deep-water reef sites on the Blake Plateau, Straits of Florida, and eastern Gulf of Mexico documenting their habitat and benthic biodiversity for the first time. We also produced a high-definition topographic SEA- BEAM map at the southwest Florida lithoherm site. The sites described here range from the Blake Plateau off South Carolina to the southwestern Florida slope and include the following: 1) Stetson Lophelia reefs: an extensive region of Lophelia bioherms and lithoherms along the eastern Blake Plateau off South Carolina, including the discovery of a 152-m tall pinnacle with thickets of REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 345

Lophelia (depth 822 m); 2) Savannah Lophelia lithoherms: numerous lithoherms with relief up to 60 m and capped with Lophelia on the western Blake Plateau off Georgia (depth 550 m); 3) east Florida Lophelia reefs: nearly 300, 15–152-m tall mounds that appear to be Lophelia bioherms and lithoherms distributed along a 222-km stretch off eastern Florida (depth 700–800 m); 4) Miami Terrace escarpment: Miocene-age terrace off southeastern Florida with high-relief, hard-bottom habitats and rich benthic communites (depth 300–600 m); 5) Pourtalès Terrace: a continuation of Miami Terrace off the Florida Keys (depth 200–460 m) of which habitat and fauna were described in detail by Reed et al. (2005a); and 6) west Florida Lophelia lithoherms: 15- m tall Lophelia lithoherms off the southwestern Florida shelf slope (depth 500 m). These sites are contrasted to the azooxanthellate deep-water Oculina reefs, which occur at depths of 70–100 m at the shelf-edge off central eastern Florida.

Methods and Materials

Human occupied submersibles and ROVs were used to conduct benthic surveys from 1999 to 2004 at six deep-water reef sites off the southeastern U.S. HBOI’s R/V Edwin Link (Seward Johnson II) and R/V Seward Johnson supported the Johnson-Sea-Link (JSL) and Clelia submersibles. These submersibles have an acrylic sphere (JSL) or hemisphere (Clelia) that provides >180° field of view to the observers within. Each was equipped with a manipulator arm with a 20-cm clam-shell grab, jaw, and suction hose; twelve 12.7-L Plexiglas buckets, and a CTD data recorder (Seabird SBE 25 Sealogger) that continuously recorded time, temperature, conductivity, salinity, oxygen, and depth. Ship navigation utilized differ- ential GPS (Magnavox MX 200 Global Positioning System), which had an estimated statistical positioning error of 1–5 m. Submersible navigation used Ultrashort Baseline Sonar (USBL) technology, which consisted of ORE Trackpoint II Acoustic Positioning System and Integrat- ed Positioning System (IPS) software that integrated the submersible’s position relative to the ship and calculated the submersible’s real time DGPS position throughout each dive. Analysis of USBL tracking accuracy for a worst-case tracking scenario estimated a maximum statistical positioning error of 9.6 m at a depth of 500 m (J. Kloske, Florida Institute of Oceanography, pers. comm.; Opderbecke, 1997). At each dive site, profiles of bottom topography were made with the ship’s echosounder (SIMRAD EQ50 video echo sounder 38/50 kHz). Color videotapes (digital mini DV or high 8-mm) were recorded during dives with an external pan and tilt videocamera (Sony DX2 3000A with Canon J8X6B KRS lens, 6–48 mm zoom, and 0.3 m minimum focus), which had parallel lasers (25 cm apart) for scale. Collected specimens were photographed in situ with an external Benthos camera with 85-mm lens (Ektachrome ASA 100) or a Nikon Coolpix 990 digital camera. The west Florida lithoherm site was surveyed in 2003 from the R/V Ronald H. Brown using the Innovator ROV (Sonsub, Inc.), an industrial ROV rated to 3000 m depth. The ROV was equipped with a manipulator arm, collection buckets, color video camera, and digital still camera. A high definition topographic map was made of the site with a SEABEAM 2112 (12 KHz) swath bathymetric sonar system capable of hydrographic charting and seafloor acoustic backscatter imaging in depths of 50–11,000 m with up to 151 beams. Video data on the east Florida Lophelia reefs were also gathered in the 1980s with the Cord ROV (HBOI). Additional information was summarized from published literature on submersible dives with Alvin (Milliman et al., 1967; Neumann et al., 1977; Griffin and Druffel, 1989; Messing et al., 1990), Aluminaut (Neumann and Ball, 1970), U.S. Navy’s NR-1 submarine (Paull et al., 2000; Weaver and Sedberry, 2001), and from surveys using echo-soundings, dredges, and camera sleds (Stetson et al., 1962; Menzies et al., 1973; Cairns, 1979; Mullins et al., 1981; Cairns, 2000). Specimens of dominant benthic macroinvertebrates (primarily sponges, cnidarians, mollusks, and echinoderms) were collected using the submersible’s manipulator. Documentation 346 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006

of specimens and habitat included in situ videotapes (100 hr), in situ photographic images (259), museum voucher specimens (621), and detailed notes, which enabled habitat characterization and description of the benthic and fish communities. Specimens, photographs, and videotapes are cataloged at HBOI’s Museum for the Division of Biomedical Marine Research (DBMR). The collections of macroinvertebrates were not quantitative, but primarily targeted the dominant, large (>5 cm), sessile organisms. Thinly encrusting organisms were not generally collected due to the difficulty of sampling these with the manipulator. These benthic taxa, for the most part, cannot be identified to species-level from videotapes, but required micro- scopic analyses of collected specimens (e.g., Porifera, Octocorallia). No fishes were collected, but were identified from videotapes and direct observations. When identifications could not be made to species level, fish were identified to genus or family levels. Qualitative estimates of size and abundances of fishes and benthic invertebrates were made visually by the observer in the submersible and from videotapes using the videocamera’s parallel lasers. Depending on the camera’s degree of zoom, the field of view in the video ranged from 25 cm to ~3 m. On most dives the submersible was kept within 1 m of the bottom while transiting from the base to the peak of the feature.

Results

For each deep-water reef region, we include a summary of the state of knowledge prior to reporting the results from our 1999–2004 expeditions. The habitat and fauna of Pourtalès Terrace was described previously in detail by Reed et al. (2005a) based on 15 submersible dives on the Terrace, including the Jordan and Marathon deep-water sinkholes and five high-relief bioherms (Fig. 1, Table 1).

Region D: Stetson Reefs, Eastern Blake Plateau This site is on the eastern edge of the Blake Plateau within the region of the Charles- ton Bump, ~220 km (120 nm) southeast of Charleston, South Carolina, at depths of 640–869 m (Table 1, Fig. 1). Thomas Stetson first described these features from echo soundings and bottom dredges as steep-sloped mounds with active coral growth on top (Stetson et al., 1962; Uchupi, 1968). Stetston estimated that over 200 coral mounds up to 146 m in height spread over a 6174-km2 area and observed live coral colonies up to 50 cm in diameter with a camera sled. Enallopsammia profunda (referred to as Dendrophyllia profunda) was the dominant species in all areas, although L. pertusa (referred to as Lophelia prolifera) was concentrated on top of the mounds. Densest coral growth occurred along an escarpment at region D. Slightly north of this region are the Agassiz coral hills (32°36′N, 76°58′W; 650 m), which consist primarily of solitary corals (Bathypsammia spp. and Flabellum spp.), but little Lophelia (George, 2002). Detailed geological maps of the region also suggest numerous potential coral mounds, but few have been confirmed by direct observation (Popenoe and Manheim, 2001). Our fathometer transects in 2002 surveyed only a fraction of the entire Stetson reef area, but revealed dozens of individual pinnacles and mounds. From these fathometer transects, two pinnacle regions were selected for submersible exploration. A small subset of the Stetson reef area was first mapped by six fathometer transects covering ~97 km2 (11.1 × 8.7 km; 31°59.03′N–32°05.03′N and 77°42.75′W–77°37.98′W), in which six major peaks or pinnacles and four major scarps were plotted. These pinnacles ranged from 689 to 643 m at their bases, with relief of 46–102 m. A subset of this was further mapped with 70 fathometer transects spaced 250 m apart (re- cording depth, latitude, and longitude every ~3 s), covering an area of 1.9 × 2.8 km REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 347

Figure 1. Deep-water coral reef regions off southeastern U.S. • = Johnson-Sea-Link I and II submersible and ROV dive sites (see Table 1 for locations); Regions: A = Oculina reefs (Reed, 1980; 2002a), B = east Florida Lophelia reefs, C = Savannah Lophelia lithoherms, D = Stetson reefs, E = Mullins reefs (Mullins et al., 1981), F = Bahama lithoherms (Neumann et al., 1977), G = Miami Ter- race escarpment, H = Pourtalès Terrace (Reed et al., 2005a), I = west Florida lithoherms.

(32°00.5′N–32°01.5′N and 77°40.0′W–77°42.5′W), and resulting in a three-dimen- sional bathymetric GIS Arcview map of a major feature, which we named “Stetson Peak.” Three JSL submersible dives were made on pinnacle 3 and four dives on Stet- son Peak, which is described below (Table 1). Stetson Peak was 780 m deep at the south base with the peak in 627 m (differen- tial GPS coordinates of submersible at the peak: 32°01.6882′N, 77°39.6648′W). This represents one of the tallest Lophelia coral lithoherms known, nearly 153 m in relief (Fig. 2). The linear distance from the south base to the peak was ~0.9 km. The lower south-facing flank of the pinnacle, from ~762 to 701 m, was a gentle slope of 10°–30° with a series of 3–4-m high ridges and terraces that were generally aligned 60°–240°. These ridges were covered with nearly 100% Lophelia coral rubble, as well as live Lophelia colonies 10–30 cm tall, and standing dead colonies 30–60 cm tall. Very little rock was exposed, except on the steeper exposed, eroded ridge faces. Some rock slabs, ~30 cm thick, slumped from these faces. From 701 to 677 m, the slope increased from ~45° to 60°. From 671 m to the peak, the geomorphology was very complex and rugged, consisting of 60°–90° 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 348 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006 (peak) (peak 1) 79°15 ′ W 31°48 ′ N, 31°41.5 ′ N, 79°08.60 ′ W 79°18.06 ′ W 79°17.46 ′ W 31°41.82 ′ N, 31°41.23 ′ N, 32°00.6302 ′ N, 79°08.5964 ′ W 79°07.4831 ′ W 79°05.5544 ′ W 79°05.2516 ′ W 77°41.9285 ′ W 77°39.6648 ′ W 31°41.4259 ′ N, 31°42.2555 ′ N, 31°44.0975 ′ N, 31°44.3814 ′ N, 32°01.6882 ′ N, (trawl records) (trawl GPS coordinates 34°26 ′ N, 75°42 W W; 34°18 ′ N, 75°47 W; 9 15 11 11 30 (m) 36.03 9 35.82 9 Salinity Vis 35.04–35.16 9 ) − 1 50 5–25 (50°) (50°) (50°) (30°) (45°) 25–40 10–15 10, 20 35–60 (45°, 30°) (60°, 30°) (direction to) (20°, 60°–75°) Current (cm s (180°, 355°, 80°) 7.5 (°C) 9.96 8.75 9.04 Temp. Temp. 8.9–9.5 7.97–8.4 25–45 8.35–8.5 6.22–8.22 5–20 10.86–10.96 10, 20 13 33 61 50 15 38 54 114 153 80–100 (2.2 nm N–S) (0.8 nm N–S) (width at base) (0.47 nm N–S) (0.53 nm N–S) Max. relief (m); (0.3 nm NE–SW) (0.4 nm NE–SW) (peak 1) Depth at peak (m) Lophelia pertusa from R/V sites #412, Combat #14449); (see R/V Terrace * Reed Eastward = region H, Pourtalès (m) 503 490 541 532549 499 488 537 487 549 533 549 511 550 500 694 579 370–450 Dive #Dive Depth at base Alvin 203Alvin 10) Savannah lithoherms10) Savannah JSL II 1,690 7) Savannah lithoherms7) Savannah JSL II 1,697 lithoherms8) Savannah JSL II 1,698 9) Site 1, pinnacle #4 JSL II 3,327 6) Site 2, pinnacle #1 JSL II 3,331; 3,332 5) Site 2, pinnacle #5 JSL II 3,330 4) Site 2, pinnacle #6 JSL II 3,328; 3,329 Region C: Savannah lithoherms Savannah C: Region site Alvin 3) *North Carolina Lophelia reefs reefs Lophelia Carolina *North (Menzies et al., 1973; Cairns, 1979; Reed and Ross, 2005) Table 1. Site summary for coral deep-water SE reefs U.S. and (in lithoherms order off N–S). Site reference numbers correspond Table numbers to were Figure not 1. part Sites of without dive * this = study. North Carolina sites (Cairns, 1979: trawl records of Alvin Alvin WHOI’s = et al., 2005a); number: ROV, dive JSL I, II Innovator = Johnson-Sea-Link = Innovator Sonsub’s I HBOI’s and II submersibles, Cord Cord = ROV, HBOI’s submersible; depth = at base, peak, maximum relief, and maximum width at base of bioherm; physical parameters are values near the maximum GPS location at peak of bioherm (or as indictated). depth; vis coordinates are submersible/ROV = estimated visibility from submersible/ROV; *Site reference Region D: Stetson reefs Stetson D: Region 1) Stetson peak2) Pinnacle #3, peaks 1–4 JSL II 3,314–3,316 JSL II 3,317; 3,320–3,322 780 627 REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 349 06.6 ′ W o (peak) (Loran C) 79°38.4 ′ W 30°48.2 ′ N, 79°41.17 ′ W 79°36.51 ′ W 28°46.72 ′ N, 28°02.04 ′ N, 79°39.4743 ′ W 79°38.9784 ′ W 79°38.0678 ′ W 79°37.5859 ′ W 79°37.6735 ′ W 79°37.0064 ′ W 79°35.3889 ′ W 79°36.8306 ′ W 79°35.5994 ′ W 79°34.9679 ′ W 30°30.1194 ′ N, 30°16.8114 ′ N, 29°40.2628 ′ N, 28°47.6258 ′ N, 28°39.8464 ′ N, 28°28.3513 ′ N, 27°01.3474 ′ N, 28°17.0616 ′ N, 27°12.5695 ′ N, 27°39.4305 ′ N, GPS coordinates coordinates GPS 28°59.2 ′ N, 80 27°32.8 ′ N, 79°56.2 W to 12 18 12 15 12 (m) 34.9 14 35.28 34.96 14 34.91 12 34.93 14 34.90 9 34.91 15 34.99, Salinity Vis ) 1 - 0 10 15 15 25 15 (5°) (5°) 5–20 (320°) (215°) (315°) (305°) (260°) (330°) (345°, 360°) (direction to) (direction Current (cm s (cm Current (340°, 360°, 50°) 6.5 7.71 7.05 10–15 7.78 6.777.24 10, 30 15–20 6.31 6.49 6.79 (°C) Temp. 8.1–8.4 5, 25, 30 7.6–8.4 7.4–26.7 0–58.5 35.7–36.4 1–30 88 61 30 91 75 44 44 24 97 42 29 46 122 150 max (0.3 nm N–S) (0.99 nm N–S) (0.53 nm N–S) (0.78 nm N–S) (0.84 nm N–S) (width at base) at (width Max. relief (m); (m); relief Max. (3 nm N–S; 0.8 E–W) (0.9 nm N–S; 0.9 E–W) (0.95 nm N–S; 0.82 nm E–W) nm 0.82 N–S; nm (0.95 (1.66 nm N–S; 1.0 nm E–W) nm 1.0 N–S; nm (1.66 (0.3 nm N–S; 0.9 nm NW–SE) nm 0.9 N–S; nm (0.3 579 Depth at at Depth peak (m) peak 866822 744 777 734 716 793 762 671 701 544 157 max; peak 6 = 107 791762 716 758 718 713 838 741 723 685 750723 721 676 (m) 70–100 (440–914) Dive # Dive base at Depth JSL I 4,659 JSL I 4,657 JSL II 3,335 (3,233–3,251) JSL II 3,333; 3,334; JSL I 4,658 JSL I 4,660 JSL I 4,661 JSL II 3,336 JSL I 4,663 JSL I 4,656 JSL I 4,662 13) Jacksonville Lophelia reef, 13) Jacksonville pinnacle #186 Augustine Lophelia reef, 14) St. pinnacle #3 Lophelia reef, 15) Cape Canaveral pinnacle #113 16) Cape Canaveral Lophelia reef16) Cape Canaveral JSL I 2,474 Region B: east Florida Lophelia reefs Lophelia Florida east B: Region (2000) lithoherm site11) Paull JSL II 3,245; Lophelia reef, 12) Jacksonville pinnacle #204B, peak 6 17) Cape Canaveral Lophelia reef, 17) Cape Canaveral pinnacle #129 Lophelia reef, 18) Cape Canaveral pinnacle #TS7 (near P 135) Lophelia reef, 19) Cape Canaveral pinnacle #151 Region A: Oculina reefs reefs Oculina A: Region (Reed, 1980; 2002a,b) 20) Cape Canaveral Lophelia reef20) Cape Canaveral 85 Cord ROV 23) Jupiter Lophelia reef, pinnacle #293 Table 1. Continued. 1. Table reference Site * 21) Ft. Pierce Lophelia reef, pinnacle #TS4 (near P 212) 22) Stuart Lophelia reef, pinnacle #292 350 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006 (base) (peak) (ridge top) GPS coordinates 27°10 ′ N, 77°30 W 27°25 ′ N, 79°25 W (base of escarpment) (base of escarpment) (base of escarpment) (base of escarpment) (base of escarpment) 27°40 ′ N, 78°15 W to 29°09.5 ′ N, 88°01.0 W 24°44.71 ′ N, 80°27.59 W 26°56.72 ′ N, 79°16.02 W to 24°15.33 ′ N, 80°54.27 W to 26°05.6902 ′ N, 79°50.2540 W 25°41.9970 ′ N, 79°51.0510 W 26°01.2885 ′ N, 79°49.3258 W 26°05.7066 ′ N, 79°50.3634 W 25°41.9959 ′ N, 79°51.8924 W 26°19.9094 ′ N, 84°45.8639 W 26°20.3915 ′ N, 84°44.8733 W 26°20.0133 ′ N, 84°45.0030 W 25°35.9864 ′ N, 79°54.2491 W 25°35.9963 ′ N, 79°52.9368 W 9 11 11 18 18 18 (m) 15–30 34.00 9 34.93 12 34.96 14 Salinity Vis 34.9–35.1 34.92–34.95 11–15 ) 1 - ) o , 180°) o 5 5 0 10 25 15 50.0 5–10 (10°) Slight Slight (360 (360°) (145°) (230°) (180°) (direction to) (direction (–, 360 Current (cm s (cm Current N/A N/A N/A (°C) 7.61 8.17 Temp. Temp. 4 40 4.0–6.0 50 8.25– 9.58 0–15 (N) 12–180 8.6–12.63 0–100.0 35.04–35.78 9–15 escarpment (width at base) Max. relief (m); Depth at peak (m) (m) 437 310 126 6.94–7.6 0, 5, 20 335 284 51 573 399 174 375 279 95 391 321 70 7.38–7.64 530 434 90 7.0–9.0 549 393 155 7.97 430 322 112 610–675 198–461 1,000–1,300 Dive #Dive Depth at base 4,667; 4,680 4,666 4,679 JSL I 4,677 Innovator ROV6Innovator ROV7Innovator 448 466 412 454 36 12 Innovator ROV 5 ROV Innovator 558 554 26) West ridge, east face, site #BU6 ridge, east face, West 26) JSL I 4,665; Region E: Mullins reefs Region (Mullins et al., 1981; Reed, 2002a) 25) East ridge, east face, site #BU425) East ridge, east face, JSL I 4,668 Table 1. Continued. 1. Table * Site reference Region F: Bahama lithoherms Bahamalithoherms F: Region (Neumann et al., Messing et 1977; al., 2002a) Reed, 1990; Terrace Miami G: Region site #BU424) East ridge, west face, JSL I 4,664; 28) West ridge, east face, site #BU2 ridge, east face, West 28) JSL I 4,670; 27) West ridge, east wall, site #BU2 ridge, east wall, West 27) JSL I 4,669 29) West ridge, base of east face, site ridge, base of east face, West 29) #BU1b Gulf of Mexico, Viosca Knolls Viosca Gulf of Mexico, 2002) (Schroeder, 32) Site 2 escarpment 33) Pinnacle #4 Region I: west Florida lithoherms Florida west I: Region 31) Pinnacle #1 Region H: Pourtalès Terrace Pourtalès H: Region 2005a) al., et (Reed (34,35) 30) West ridge, west face, site #BU1b ridge, west face, West 30) JSL I 4,678 REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 351

Figure 2. Echosounder profiles of deep-water reefs off southeastern U.S. A) Stetson Peak (780 m base, 627 m peak); B) Savannah lithoherm, site 2, pinnacle 1 (537 m base, 487 m peak); C) east Florida Lophelia pinnacle 204B (701 m base, 544 m peak); D) east Florida Lophelia pinnacle 113 (777 m base, 716 m peak); E) Miami Terrace escarpment, site BU1b (549 m base, 322 m peak); F) Pourtalès Terrace, Tennessee bioherm #2 (213 m base, 131 m peak), g) SEABEAM bathymetry of escarpment and lithoherms at west Florida lithoherm site (depth 412 m at top of escarpment, 500 m on lithoherm plateau; vertical exaggeration 2.5×). thick rock crust, which was undercut from erosion; below this was a 90° escarpment of 3–6 m. The peak was a flat rock plateau at 625–628 m and was ~190 m across on a S-N submersible transect. The north face was not explored in detail, although a vertical rock wall from the peak to ~654 m graded downward to a 45° slope with boulders and rock outcrops. 352 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006

Table 2. Species list of macroinvertebrates associated with deep-water reefs off SE U.S. (Sites: SR = Stetson reefs; SL = Savannah lithoherms; FL = Florida east coast Lophelia reefs; MT = Miami Terrace escarpment; PT = Pourtalès Terrace; WF = west Florida lithoherms).

Taxonomy Max depth Min depth SR SL FL MT PT WF (m) (m) Phylum PORIFERA PORIFERA, Class DEMOSPONGIAE Aka (Siphonodictyon) sp. 648 183 X XX Ancorina? sp. 641 641 X Ancorinidae?, unid. sp. 586 586 X Astrophorida, unid. sp. 509 173 X X X Astrophorida, new sp.? 520 520 X Auletta sp. 207 171 X Axinellida, unid. sp. 499 168 X X Axinellida + Plakortis? sp. 210 210 X Biemnidae, unid. sp. 628 512 X X Callipelta? sp. 206 206 X Calthropellidae, unid. sp. 757 757 X Characella? sp. 198 198 X Chondrosia? sp. 300 297 X Corallistes sp. 689 226 X X Corallistidae, unid. sp. 767 186 X X X Demospongiae, unid. sp. 541 170 XX Dercitus cf. bucklandi (Bowerbank, 1858) 809 809 X Dictyoceratida, unid. sp. 222 172 X Discodermia sp. 269 180 X Echinodictyum sp. 172 171 X Epipolasis sp. 211 211 X Erylus sp. 356 216 XX Erylus transiens (Weltner, 1882) 262 262 X Geodia sp. 767 174 X X X Geodiidae, unid. sp. 816 180 XX Halichondrida, unid. sp. 260 260 X Halichondriidae, unid. sp. 648 237 X X Haplosclerida, unid. sp. 543 171 X Hymedesmia sp. 1 (blue morph) 179 172 X Hymedesmia sp. 2 (yellow morph) 179 172 X Ircinia new sp.? 500 500 X Leiodermatium sp. 754 172 X X X Lithistida, unid. sp. 310 185 X Mycalidae, unid. sp. 312 284 X Oceanapia sp. 652 172 X X Oceanapiidae, unid. sp. 758 758 X Pachastrellidae, unid. sp. 811 166 X X X X X Petrosina?, uind. sp. 183 183 X Petrosiidae, unid. sp. 750 178 X X X X X Phakellia new sp. 1 171 171 X Phakellia new sp. 2 174 174 X Phakellia new sp. 3 174 174 X Phakellia spp. 756 171 X X X X X X REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 353

Table 2. Continued.

Taxonomy Max depth Min depth SR SL FL MT PT WF (m) (m) Plakinidae, unid. sp. 660 638 X Plakortis sp. 312 210 X Poecillastra? sp. 427 323 XX Poecilosclerida, unid. sp. 717 132 XX Polymastia sp. 726 726 X Raspailiidae, unid. sp. 763 321 X X X X Spirophorida, unid. sp. 183 183 X Spongosorites sp. 671 171 X X X X Stellettidae?, unid. sp. 312 312 X Stellettinopsis? sp. 198 198 X Stylocordyla sp. 515 515 X Theonellidae, unid. sp. 472 470 X Theonellidae, new genus, new sp. 208 199 X Topsentia? sp. 173 173 X Vetulina? sp. 415 415 X Zyzzya sp. 222 222 X PORIFERA, Class HEXACTINELLIDA Aphrocallistes sp. 800 587 X Heterotella sp. 762 418 X X X Hexactinellida, unid. sp. 800 186 X X X X X X Hexasterophora, unid. sp. 761 517 X X X Hyalonematidae?, unid. sp. + Zoanthidae 737 737 X Lychniscosida, unid. sp. 662 649 X Lyssacinosida, unid. sp. 757 628 X X Phylum CNIDARIA CNIDARIA, Class HYDROZOA Hydroida, unid. sp. 656 202 X X Hydroida, unid. sp.1 322 284 X HYDROZOA, STYLASTERIDAE Distichopora foliacea Pourtalès, 1868 175 175 X Pliobrothus echinatus Cairns, 1986 175 175 X Stylaster erubescens Pourtalès, 1868 186 175 X Stylaster filogranus Pourtalès, 1871 200 175 X Stylaster miniatus (Pourtalès, 1868) 200 175 X Stylasteridae, unid. sp. 742 173 X X X X X CNIDARIA, Class ANTHOZOA ANTHOZOA, ACTINIARIA Actiniaria, unid. sp. 751 565 X Hormathiidae (Venus fly trap anemone) 750 284 XX ANTHOZOA, ANTIPATHARIA Antipatharia, unid. sp.1 (re-or morph) 767 283 X X X X Antipatharia, unid. sp.2 (wh-pi morph) 515 328 X X X X Antipathes rigida? Pourtalès, 1868 319 319 X Bathypathes alternata Brook, 1889 716 466 XX ANTHOZOA, SCLERACTINIA Bathypsammia? sp. 640 418 X X Enallopsammia profunda (Pourtalès, 1867) 742 305 X X X X Lophelia pertusa (Linnaeus, 1758) 815 284 X X X X X Madrepora oculata Linnaeus, 1758 763 288 X X X 354 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006

Table 2. Continued.

Taxonomy Max depth Min depth SR SL FL MT PT WF (m) (m) Scleractinia, unid. sp. 632 582 X X Solenosmilia variabilis Duncan, 1873 470 470 X ANTHOZOA, OCTOCORALLIA Anthomastus cf. agassizzi Verrill, 1922 757 420 XX Candidella imbricata (Johnson, 1862) +Thouarella? sp. 732 732 X Capnella nigra (Pourtalès, 1868) 768 325 X X X Chrysogorgia squamata (Verrill, 1883) 581 581 X Clavularia new sp.? 648 648 X Echinomuricea cf. atlantica (Johnson, 1862) 323 284 X Eunicella modesta (Verrill, 1883) 732 518 X X Isidella sp. 1 762 744 X Isididae, unid. sp. (bamboo coral) 816 340 X X X X Keratoisis flexibilis (Pourtalès, 1868) (pink morph) 734 374 XX Keratoisis flexibilis (Pourtalès, 1868) (white morph) 816 378 X X X X Muriceides hirta? (=Trachymuricea) (Pourtalès, 1867) 716 681 X Muriceides sp. (not hirta, not kukenthali) 191 191 X Paramuricea cf. echinata Deichmann, 1936 716 716 X Paramuricea multispina Deichmann, 1936 715 189 XX Paramuricea cf. multispina Deichmann, 1936 323 323 X Paramuricea placomus (Linnaeus, 1758) 470 462 X Paramuricea cf. placomus (Linnaeus, 1758) 326 283 X Paramuricea unid. sp. 762 322 X X Placogorgia mirabilis Deichmann, 1936 212 172 X Placogorgia tenuis? (Verrill, 1883) 557 457 X Placogorgia? sp. 1 579 565 X Plexauridae, unid. sp. 1 716 579 X X Plumarella pourtalesii (Verrill, 1883) 753 171 X X X X X Pterostenella? new sp.? 754 754 X Swiftia casta (Verrill, 1883) 525 525 X Swiftia new sp.? 497 497 X Telestula? sp. 2 784 734 X Thesea parviflora Deichmann, 1936 183 183 X Thouarella? sp. 732 732 X Trachymuricea hirta (Pourtalès, 1867) 468 462 X Villogorgia cf. nigrescens Duchassaing & Michelotti, 1860 215 215 X ANTHOZOA, ZOANTHIDEA Zoanthidae, unid. sp.1 (orange morph) 656 519 X X Zoanthidae, unid. sp.1 (yellow morph) 649 502 X X Zoanthidae, unid. sp.2 734 734 X Zoanthidae, unid. sp.3 699 419 XX Zoanthidae, unid. sp.4 328 328 X Phylum ARTHROPODA, DECAPODA Bathynectes longispina Stimpson, 1871 (bathyal swimming crab) 662 514 X X X Bathynomus giganteus A. Milne Edwards, 1879 (giant isopod) 396 396 X Chaceon fenneri (Manning & Holthuis, 1984) (golden crab) 766 455 X X X X X X Eumunida picta Smith, 1883 (squat lobster) 714 285 X X X X X REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 355

Table 2. Continued.

Taxonomy Max depth Min depth SR SL FL MT PT WF (m) (m) Phylum MOLLUSCA Bursa tenuisculpta (Dautzenberg & Fischer, 1906) 283 187 XX Calliostoma pulchrum (C.B. Adams, 1850) 187 187 X Conus villepini Fisher & Bernardi, 1857 188 171 X Entemnotrochus adansonianus (Crosse & Fischer, 1861) 265 180 X Hyalina albolineata (Orbigny, 1842) 187 187 X Megalocranchia cf. oceanica (Voss, 1960) 729 729 X Murex beauii Fischer & Bernardi, 1857 188 188 X Perotrochus amabilis (F.M. Bayer, 1963) 265 181 X Perotrochus midas F.M. Bayer, 1965 393 262 X Scaphella gouldiana (Dall, 1887) 188 187 X Phylum BRYOZOA Membranipora? sp. 631 631 X Phylum ECHINODERMATA Asteroidea, unid. sp. 653 454 X X X Crinoidea, Comatulida, unid. sp. 751 373 X X X Holothuroidea, unid. sp. 181 181 X Ophiuroidea, Gorgocephalidae, unid. sp. 304 304 X

Dominant sessile macrofauna consisted of Scleractinia, Stylasteridae, Octocoral- lia, and Porifera (Table 2). The colonial Scleractinia were dominated by colonies of L. pertusa (30–60 cm tall) accompanied by E. profunda and S. variabilis. Small stylasterid corals (15 cm tall) were common and numerous species of solitary cup corals were abundant. Dominant Octocorallia (six taxa total) consisted of Prim- noidae (15–30 cm tall), Paramuriceidae (60–90 cm), Isididae bamboo coral (15–60 cm), Stolonifera, and Nephtheidae (5–10 cm). Eighteen sponge taxa were identified. The most abundant taxa were Pachastrellidae (25-cm fingers and 25–50-cm plates), Corallistidae (10-cm cups), Hexactinellida glass sponges (30-cm vases), Geodia spp. (15–50-cm spheres), and Leiodermatium spp. (50-cm frilly plates). On the steeper upper flank, from 671 to 625 m the density, diversity, and size of sponges increased, with 15–50 cm sponges more abundant. Massive Spongosorites spp. and Hexactinel- lida were common, and Pachastrellidae tube sponges were abundant. On the peak plateau, the dominant macrofauna consisted of colonies of L. pertusa (30–60 cm tall), Phakellia spp. fan sponges (30–50 cm), and numerous other demosponges. Coral rubble was also abundant. Although motile benthic invertebrates were not targeted for collection, some dominant groups were noted. Squat lobsters Eumunida picta (Galatheidae), and the crabs Bathynectes longispina (Portunidae) and Chaceon fenneri (Geryonidae) were the most common decapod crustaceans. Two species of echinoids were common, Stylocidaris spp. and an unidentified white urchin. No ho- lothurians 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. Ten species of fish were identified from videotapes, primarily red bream (Beryx decdactylus), roughy (Hoplostethus occidentalis), codling (Laemonema melanurum), rattail (Nezumia spp.), and hake (Phycidae) (Table 3). No large fish were seen on the plateau. 356 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006 X X X X X X Lophelia X XX X XXX XX X X X X X X X X X X X X X SR SL FL OR MT PT WF (m) Min depth (m) 671469100 287100 322 X 70 70 100628296 70 526100 296 519 X 381198 70 296 381 171 X X 100 70 522223100100 335 100 172 70 70 70 100283415 70 179 415 100 70 Spotfin hogfish Black sea bass Common name Max depth Sheepshead ApogonidaeRed bream Labridae Ophidiidae cardinalfish Twospot Carangidae Alphonsino Carangidae 100 Brotula Blue runner jack Crevalle 70 Serranidae ChaetodontidaeChaetodontidaeChaunacidaebutterflyfish Spotfin Chlorophthalmidae Reef butterflyfish Greeneye Sea toads Pomacentridae Congridae Congridae Purple reeffish 100 70 Priacanthidae Conger eel Conger eel Bulleye Family Caproidae Deepbody boarfish 219 174 ScombridaeSerranidae Serranidae Wahoo bass Yellowfin Streamer basses Chlorophthalmidae, unid. sp. Congridae, unid. sp. Longley, 1932 pseudomaculatus Longley, Apogon 1829 Beryx decadactylus Cuvier, 1860) (Poey, Bodianus pulchellus Brotulidae, unid. sp. crysos (Mitchill, 1815) Caranx hippos (Linnaeus, 1766) Caranx falciformis (Müller & Henle, 1839) Carcharhinus Carcharhinidae Goode & Bean, 1878Caulolatilus microps 1887) ocyurus (Jordan & Evermann, Centropristis Serranidae philadelphica (Linnaeus, 1758)Centropristis shark Silky striata (Linnaeus, 1758) Centropristis MalacanthidaeChaetodon ocellatus Bloch, 1787 Serranidae 1860 Chaetodon sedentarius Poey, Bank sea bass Blueline tilefish Chaunax sp. Bonaparte, 1840 agassizi Chlorophthalmus Rock sea bass Jordan & Gilbert, 1882 enchrysura Chromis 1968 scotti Emery, Chromis oceanicus (Mitchell, 1818) Conger Chlorophthalmidae Pomacentridae Shortnose greeneye 1829) japonicus (Cuvier, Cookeolus reeffish Yellowtail 522 100 396 70 reefs; MT = Miami Terrace escarpment; PT = Pourtalès Terrace; WF = west Florida lithoherms). Terrace; escarpment; PT = Pourtalès Terrace reefs; OR = Oculina MT Miami Taxonomy 1843 Lowe, Antigonia capros Table 3. Species list of fish associated with deep-water reefs off SE U.S. (Sites: SR = Stetson reefs; SL = Savannah lithoherms; FL = Florida east coast east Florida = FL lithoherms; Savannah = SL reefs; Stetson = SR (Sites: U.S. SE off reefs deep-water with associated fish of list Species 3. Table (Cuvier, 1832) Acanthocybium solandri (Cuvier, 1792) (Walbaum, probatocephalus Archosargus Firth, 1933 Anthias nicholsi Sparidae Anthiinae REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 357 X X X X X XX X X X X X X X XX XX X X X X X X X X X XX X X 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 100 466 376 100 100 100 100 100 198 308 287 287 100 550100 550 X 100 518 518 518 392 100 100 179 179 100 100 100 100 194 174 754191 179 X X X X X 100 100 Common name Max depth (m) Min depth (m) SR SL FL OR MT PT WF Red dory Red grouper Greenband wrasse Slippery dick Painted wrasse Painted Carangidae Round scad Family Holocentridae soldierfish Spinycheek 100 Zeidae Serranidae grouper Warsaw Sciaenidae Jacknife fish Etmopteridae Lantern shark Scyliorhinidae Scyliorhinidae Marbled catshark Muraenidae Goldentail moray Haemulidae Tomtate Labridae Labridae Serranidae Streamer bass Pomacanthidae Queen angelfish (Cuvier, 1829) Decapterus punctatus (Cuvier, Diplectrum formosum (Linnaeus, 1766) Serranidae Sand perch Table 3. Continued. Table Taxonomy Agassiz, 1831 Agassiz, spinosus Corniger (Lowe, 1843) (Lowe, Cyttopsis rosea Epinephelus adscensionis (Osbeck, 1765) Serranidae Rock hind Epinephelus drummondhayi Goode & Bean, 1878 Serranidae Speckled hind Epinephelus itajara (Lichtenstein, 1822)Epinephelus itajara Serranidae Goliath grouper (Valenciennes, 1828)Epinephelus morio (Valenciennes, Serranidae Epinephelus nigritus (Holbrook, 1855) (Valenciennes, 1828)Epinephelus niveatus (Valenciennes, Serranidae grouper Snowy (Poey, 1852)) mystacinus (Poey, Epinephelus sp. ( cf. Serranidae Misty grouper? Euthynnus alletteratus (Rafinesque, 1810)Euthynnus alletteratus Scombridae Little tunny Equetus lanceolatus (Linnaeus, 1758) Etmopterus sp. (Péron & Lesueur, 1822) cuvieri (Péron & Lesueur, Galeocerdo Carcharhinidae shark Tiger (Nichols, 1927) Galeus arae Gephyroberyx darwinii (Johnson, 1866)Gephyroberyx Trachichthyidae Big roughy Gymnothorax miliaris (Kaup, 1856) Gymnothorax Gymnothorax nigromarginatus (Girard, 1858) nigromarginatus Gymnothorax Muraenidae moray Blackedge Gymnothorax sp. (cf. funebris Ranzani, 1840)Gymnothorax Muraenidae Unid. moray eel Cuvier, 1830 Cuvier, Haemulon aurolineatum (Beebe & Tee-Van, 1932)Tee-Van, bathyphilus (Beebe & Halichoeres Labridae Halichoeres bivittatus (Bloch, 1791) Halichoeres (Poey, 1860) caudalis (Poey, Halichoeres Helicolenus dactylopterus (Delaroche, 1809)Hemanthias sp. 1885)Hemanthias vivanus (Jordan & Swain, Scorpaenidae Serranidae Blackbelly rosefish Red barbier Holacanthus bermudensis Goode, 1876 Pomacanthidae Blue angelfish Holacanthus ciliaris (Linnaeus, 1758) 358 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006 X XX X X X X X X X X X XX X X X X X X X X X XX X X 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 100 100 752100 322 X X X X 586369500757 586 369 296 628 X X 100 100 100 180 100 100 100 100 100 750100 750 461650496762 461 287 650 189 770 714 X 100 284 186 X X X X X 100 Sparidae Red porgy Macrouridae Batrachoididae Grenadier Leopard toadfish Triakidae Triakidae Myctophidae Dogfish Myxinidae Dogfish Lanternfish Scorpaenidae Hagfish scorpionfish Spinycheek 100 Serranidae Scamp grouper Serranidae Black grouper Molidae Ocean sunfish Mobulidae Mobulidae Giant manta Gobiidae goby Bluegold Gobiidae Island goby Lutjanidae Lane snapper Lutjanidae Gray snapper Lophiidae Lutjanidae Goosefish Red snapper Trachichthyidae roughy Silver Roughy Trachichthyidae Trachichthyidae Roughy Chimaeridae Centrolophidae Spotted ratfish Barrelfish Moridae Moridae Shortbeard codling Codlng 521 521 X FamilyHolocentridae Squirrelfish Common name Max depth (m) Min depth (m) SR SL FL ORMT PT WF (Bloch & Schneider, 1801) acuminatus (Bloch & Schneider, Pareques Sciaenidae High-hat (Linnaeus, 1758) pagrus Pagrus N. bairdii, N. aequalis, or atlantica) Nezumia sp . (3 spp. N. bairdii, (Goode & Bean, 1880) Opsanus pardus Triakidae Mustelus sp. Myctophidae Myxine glutinosa Linnaeus, 1758 1935 Neomarinthe hemingwayi Fowler, Jordan & Swain, 1884 phenax Jordan & Swain, Mycteroperca Mycteroperca microlepis (Goode & Bean, 1879) microlepis Mycteroperca Serranidae Gag grouper (Poey, 1860) bonaci (Poey, Mycteroperca Mola mola (Linnaeus, 1758) (Walbaum, 1792) (Walbaum, Manta birostris Böhlke & Robins, 1960 spilus Böhlke Lythrypnus Böhlke & Robins, 1960 nesiotes Böhlke Lythrypnus Lutjanus synagris (Linnaeus, 1758) Lutjanus synagris Lutjanus griseus (Linnaeus, 1758) Lophius sp. 1860) (Poey, Lutjanus campechanus Cuvier, 1829 Cuvier, Hoplostethus mediterraneus 1973 Hoplostethus occidentalis Woods, Hoplostethus sp. sp. Hydrolagus (Mitchill, 1818) perciformis Hyperoglyphe Laemonema barbatulum Goode & Bean, 1883 Laemonema melanurum Goode & Bean, 1896 1962) eukrines (Starck & Courtenay, Liopropoma Serranidae Wrasse basslet Holocentrus adscensionis (Osbeck, 1765) Table 3. Continued. Table Taxonomy REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 359 X X X X X X XX X X X X XX XX X X X X X X X X XXXX XX 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 70 438 557 X 413 283 339 X X X X 186326 517 X X X X X 100 100 438 767 172 414 693 100 100 100 179 100 212 738 100 100 100 100 100 100 100 752 326 517 187 179 Common name Max depth (m) Min depth (m) SR SL FL ORMT PT WF Skate Family Peristediidae Phycidae Armored searobin Hake Polymixidae Beardfish Pomacanthidae Priacanthidae French angelfish Priacanthidae Chaetodontidae Bigeye Short bigeye Bank butterflyfish Rajidae Serranidae Spotted soapfish Clupeidae Spanish sardine Scorpaenidae Scorpaenidae Barbfish Scyliorhinidae? Scyliorhinidae Scorpionfish Carangidae Catshark Carangidae Chain digfish Greater amberjack Almaco jack Miller & Woods, 1988Woods, iwamotoi Miller & Pareques Sciaenidae Blackbar drum Table 3. Continued. Table Taxonomy (Jordan & Eigenmann, 1889) umbrosus Pareques Sciaenidae Cubbyu Peristedion sp. Peristedion Phycis sp. garrupellus Robins & Starck, 1961Plectranthias Serranidae Apricot bass Polymixia sp. Polymixia 1801) americanus (Bloch & Schneider, Polyprion Polyprionidae Wreckfish Pomacanthus arcuatus (Linnaeus, 1758) arcuatus Pomacanthus Pomacanthidae Gray angelfish Pomacanthus paru (Bloch, 1787) Pomacanthus 1829 Cuvier, Priacanthus arenatus alta (Gill, 1862) Pristigenys aya (Jordan, 1886) Prognathodes martinicensis (Guichenot, 1868)Pronotogrammus Serranidae Roughtongue bass Raja sp. 1829) (Cuvier, Rhomboplites aurorubens Lutjanidae snapper Vermilion Rypticus maculatus Holbrook, 1855 1801)Rypticus saponaceus (Bloch & Schneider, Serranidae Greater soapfish Valenciennes, 1847 aurita Valenciennes, Sardinella 1829) cavalla (Cuvier, Scomberomorus Scombridae King mackeral Scomberomorus maculatus (Mitchill, 1815)Scomberomorus Scombridae Spanish mackeral Cuvier, 1829 Cuvier, Scorpaena brasiliensis & Hildebrand, 1940Scorpaena dispar Longley Scorpaenidae Scorpaenidae Scyliorhinidae? (Garman, 1881) Scyliorhinus retifer Hunchback scorpionfishSeriola dumerili (Risso, 1810) 1833 Seriola rivoliana Valenciennes, 100 360 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006 X XX X X X X X 70 70 70 70 518 X 297 530 X 632 X X 501 X 507 X 322 518 297 100 770 529 100 100 525 399 Common name Max depth (m) Min depth (m) SR SL FL ORMT PT WF Xiphiidae Swordfish Phycidae Phycidae hake Phycid Torpedinidae Torpedinidae Atlantic torpedo ray 530 Pomacentridae Cocoa damselfish Synaphobranchidae? Cutthroat eel Squalidae Dogfish Serranidae Tattler Serranidae Belted sandfish Squalidae Dogfish Family Squalidae Dogfish Xiphias gladius Linnaeus, 1758 sp. Urophycis Torpedo nobiliana Bonaparte, 1835 Torpedo Stegastes variabilis (Castelnau, 1855) Stegastes Synaphobranchidae? Squalus sp. Poey, 1851 phoebe Poey, Serranus Serranus subligarius (Cope, 1870) Serranus (Griffith & Smith, 1834) (Griffith Sphyrna lewini 1936 Rivero, Squalus cubensis Howell Sphyrnidae Scalloped hammerhead 100 Table 3. Continued. Table Taxonomy Squalidae REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 361

Region C: Savannah Lithoherms, Western Blake Plateau A number of high-relief lithoherms occur within this region of the Blake Plateau, ~167 km (90 nmi) east of Savannah, Georgia (Table 1; Fig. 1). 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. 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°–37° 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. 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). Our fathometer transects in 2002 at Savannah lithoherm site 1 extended 4.4 km S–N (31°40.3898′N–31°42.7558′N along 79°08.5′W) and 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. The individual pinnacles ranged from 9 to 61 m in height. A single S–N submersible transect on pinnacle 4 showed a minimum depth of 499 m. The south flank of the pinnacle was a 10°–20° slope, with ~90% cover of coarse sand, coral rubble, and some 15-cm tall rock ledges. The peak was a sharp ridge, oriented NW–SE, per- pendicular to the prevailing 50 cm s−1 (1 kn) current. The north face of the ridge was a 45°, ~3-m tall rock escarpment that dropped onto a flatter terrace. From a depth of 499–527 m, the north slope formed a series of terraces or shallow depressions, ~9–15 m wide, separated by 3-m high escarpments of 30°–45°. Exposed rock surfaces were black phosphoritic pavements. The dominant sessile macrofauna occurred on the exposed terrace pavements, and greatest abundance was along the rock outcrop edges and on the pinnacle crest. The estimated cover of sponges and gorgonians was 10% on exposed rock areas. Colonies of L. pertusa (15–30 cm diameter) were common, but only accounted for ~1% of coverage. Dominant Cnidaria included several species of Octocorallia (15–20 cm tall; Primnoidae, Plexauridae), Antipathes spp. (1 m tall), and L. pertusa (Table 2). Dominant sponges included large Phakellia ventilabrum fan sponges (30–90 cm diameter), Pachastrellidae plate sponges (30 cm), Astrophorida plate sponges (30 cm), and Hexactinellida glass sponges. Our fathometer transect at Savannah lithoherm site 2 extended 8.5 km SW–NE (31°42.0812′N, 79°07.6333′W–31°45.5025′N, 79°04.0797′W) and mapped eight pinnacles with a maximum depth of 549 m and relief of 15–50 m (Fig. 10). 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 its peak was 487 m (Fig. 2). The south face, from a depth of 518–510 m, was a 10° slope, covered with coarse brown sand and Lophelia rubble. A 3-m high ridge of phosphoritic rock, extending NE–SW, cropped out at a depth of 510 m. This was nearly 100% covered by 15-cm thick standing dead Lophelia and abundant live colonies (15–40 cm). A series of rock ridges and terraces 3–9 m tall with 45° slopes were exposed between 500 and 495 m. Some of the terraces were ~30 m wide and each ridge and terrace had thick layers of standing dead Lophelia and dense live coral. Some ridges and terraces had nearly 100% cover of sponges (Phakellia spp., Geodia spp., Pachastrellidae, and Hexactinellida), Scler- actinia (L. pertusa, M. oculata), stylasterid hydrocorals, numerous species of Oc- 362 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006

tocorallia (Isididae, Primnoidae), and 1-m tall black coral bushes (Antipathes spp.). Thick deposits of sand and coral rubble filled in the depressions between the ridges. The north face was a 10° slope from 500 to 524 m covered in thick deposits of coarse brown foraminiferal sand and coral rubble. Exposed rock pavements were sparse on the north slope, but a few low rises with live bottom habitat occurred at 524 m. Dominant motile fauna at region C included decapod crustaceans (C. fenneri and Galatheidae) and a few mollusks. Ten species of fish were observed including swordfish (Xiphias gladius, 1.5 m) and sharks (Squalus spp., 60 cm). Rattail fish (Macrou- ridae) and blackbelly rosefish (Helicolenus dactylopterus) were common (Table 3). Wenner and Barans (2001) noted that wreckfish P ( olyprion americanus) were fre- quently seen on these reefs, but we saw none.

Region B: East Florida Lophelia Reefs Numerous high-relief Lophelia reefs occur in this region at the base of the Florida- Hatteras Slope at depths of 670–866 m (Table 1, Fig. 1). The reefs in the southern portion of this region lie along the western edge of the northern Straits of Florida, 28–46 km east of the Oculina coral reefs marine protected area (MPA). In 1982, dives with Cord ROV first confirmed that some of these mounds were in fact Lophelia reefs (Reed, 2002b). The northern sites off Jacksonville and southern Georgia are lithoherms that are rocky pinnacles capped with coral debris and live coral thickets (described in part by Paull et al., 2000), whereas the features from south of St. Augustine to Jupiter appear to be predominately unlithified sediment mounds capped with dense 1-m tall thickets of L. pertusa and E. profunda with varying amounts of coral debris and live coral. In 2002 and 2004, using a single beam echosounder, we mapped nearly 300 mounds from 8 to 168 m in height along a 222-km stretch off northeastern and central Florida (from Jacksonville to Jupiter; Fig. 3). At the northern end of this region, the echosounder transects revealed a massive lithoherm, 5.7-km long (N–S), which consisted of at least seven individual peaks with heights of 30–60 m (Fig. 2). The maximum depth was 701 m with total relief of 157 m. Three submersible dives (JSL II-3333, -3334; JSL I-4658) were made on peak 6 of pinnacle 204B (30°30.1194′N, 79°39.4743′W). This was the tallest individual feature of the lithoherm and had a maximum relief of 107 m and a minimum depth at the peak of 544 m. The east face was a 20°–30° slope that became steeper (50°) near the top. The west face was a 25°– 30° slope, which steepened to 80° from 561 m to the top ridge. The slopes consisted of sand and mud, rock pavement, and rubble. A transect up the south slope revealed a 30°–40° slope with a series of terraces and dense thickets of dead and live Lophelia coral (30–60 cm tall) mostly restricted to the tops of mounds, and along terrace edges and ridges. One peak at 565 m had dense thickets of live and dead standing Loph- elia (~20% live) and outcrops of coral rubble. Dominant sessile fauna consisted of L. pertusa, abundant Isididae (30–60 cm), Antipatharia, and abundant small octocorals including the gorgonians, Placogorgia spp., Chrysogorgia spp., and Plexauridae, and Nephtheidae soft corals (Anthomastus spp., Capnella spp.). Dominant sponges consisted of Geodia spp., Phakellia spp., Spongosorites spp., Petrosiidae, Pachastrellidae, and Hexactinellida (Table 2). Farther south, off Cape Canaveral, echosounder transects on Lophelia pinnacle 113 (28°47.6258′N, 79°37.5859′W) revealed a 61-m tall pinnacle with a maximum depth of 777 m (Table 1; Figs. 1, 2). The width (NW–SE) was 1.7 km and consisted of REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 363

Figure 3. Heights of 300 mounds from echosounder transects from Jacksonville to Jupiter (left– right), Florida, at depths of 600–800 m. Submersible and ROV surveys verified that many of these are Lophelia bioherms and lithoherms.

at least three individual peaks or ridges on top, each with 15–19 m relief. One submersible dive (JSL II-3335) reported 30°–60° slopes, with sand, coral rubble, and up to 10% live coral cover. No exposed rock was observed. This appeared to be a classic Lophelia sediment mound (Mullins et al., 1981). The second dive site (JSL II-3336) at pinnacle 151 (28°17.0616′N, 79°36.8306′W) was also a Lophelia mound ~0.3 nm wide (N–S) comprised entirely of coral and sediment (Table 1). Maximum depth was 758 m, with 44 m relief. The top was a series of ridged peaks from 713 to 722 m in depth. The lower flank of the south face was a 10°–20° slope of fine light colored sand with a series of 1–3-m high linear sand dunes or ridges oriented NW–SE. The ridges had ~50% cover of L. pertusa thickets consisting chiefly of dead, standing, and intact colonies (1 m tall). Approximately 1%–10% were alive on the outer 15–30 cm of the standing dead bases. There was very little broken dead coral rubble in the sand, and we saw 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 high sand waves. The upper slope steepened to 45° and then 70°–80° near the uppermost 10 m. The top of the pinnacle had up to 100% cover of 1–1.5-m tall coral thickets on a narrow 5–10-m wide ridge. Here, the coral consisted of both L. pertusa and E. profunda. Approximately 10%–20% cover was live coral (30–90 cm tall). The north slope was nearly vertical (70°–80°) for the upper 10 m with a series of coral thickets on terraces and ridges below. No exposed rock was visible and the entire pinnacle appeared to be a Lophelia sediment mound. Corals consisted of L. pertusa, E. profunda, M. oculata, and some Stylasteridae. Dominant Octocorallia included Primnoidae (two species), Isididae (Isidella spp. and Keratoisis flexibilis), and the alcyonaceans Anthomastus spp. and Capnella spp. (Table 2, Fig. 4). Dominant sponges consisted of several species of Hexactinellida, large yellow demosponges (60–90 cm diameter), Phakellia spp., and Pachastrellidae. In total, 30 Cnidaria and 18 Porifera taxa were collected from these sites. Motile fauna consisted primarily of echinoderms: cidaroid and Hygrosoma spp. urchins, co- matulid crinoids, but no stalked crinoids. Some large decapod crustaceans included C. fenneri and large Galatheidae. No mollusks were observed but were likely within the coral habitat, which was not collected. Twelve species of fish were identified and 364 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006

Figure 4. Deep-water reef habitat and associated sessile fauna off southeastern U.S. (A = Para- muricea placomus octocoral with ophiuroids, Miami Terrace escarpment, 304 m; B = Keratoisis flexibilis bamboo coral, east Florida Lophelia reefs, 592 m; C = Lophelia pertusa with Telestula sp. octocoral, east Florida Lophelia reefs, 734 m; D = Phakellia sp. sponge, Stetson reefs, 622 m; E = Hormathiidae Venus fly trap anemone, east Florida Lophelia reefs, 596 m; F = Petrosiidae sponge, Miami Terrace escarpment, 373 m).

included blackbelly rosefish H( . dactyopterus), chimaera (Hydrolagus spp.), codling (L. melanurum), goosefish Lophius ( sp.), dogfish (Mustelus spp.), hagfish (Myxine glutinosa), rattail (Nezumia spp.), cutthroat eel (Synaphobranchidae), and wreckfish ( P. americanus) (Table 3).

Region G: Miami Terrace Escarpment 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 (Kofoed and Malloy, 1965; Uchupi, 1966; Uchupi and REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 365

Figure 5. Deep-water reef habitat and associated sessile fauna off southeastern U.S. (A = Lophelia pertusa coral thicket, west Florida lithoherms, 457 m; B = antipatharian black coral with Stylaster spp., Miami Terrace escarpment, 283 m; C = Heterotella sp. vase sponge and Pachastrellidae plate sponge, Savannah lithoherms, 512 m; D = Biemnidae sponge in Lophelia coral thicket, Savannah lithoherms, 512 m; E = Lophelia pertusa and Echinomuricea cf. atlantica octocoral, Miami Terrace escarpment, 284 m; F = Petrosiidae sponge, Stetson reef, 647 m).

Emery, 1967; Uchupi, 1969; Malloy and Hurley, 1970). Previous investigations include geological studies (Neumann and Ball, 1970; Ballard and Uchupi, 1971) and dredge- and trawl-based faunal surveys in the 1970s (e.g., Halpern, 1970; Holthuis, 1971, 1974; Cairns, 1979). Lophelia mounds are 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 Madrepora growing on elongate depressions, sand ridges, and mounds at the base of the Miami Terrace escarpment. Large quantities of L. pertusa and E. profunda have also been dredged from 738 to 761 m at 26°22′N–26°24′N and 79°35′W–79°37′W (Cairns, 1979). Our JSL dives and echosounder transects in 2004 at four sites along the eastern edge of the Miami Terrace (sites BU1b, 2, 4, and 6) revealed that the outer Terrace rim consisted of a double (N–S) ridge system with steep, phosphoritic limestone escarp- 366 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006

ments of apparent Miocene age and relief up to 90 m. The ridges were capped withL. pertusa, Stylasteridae, Isididae, and various sponges and octocorals (Table 1, Fig. 2). At site BU4, the narrow N–S trending east ridge was 279 m at the top and had a steep 95-m tall western escarpment. The east and west faces of the ridges were 30°–40° 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 in 310 m, and the base of the valley between the west and east ridges was in 420 m. At site BU2, the echosounder transect showed a 13-m tall, rounded mound in 636 m near the base of the terrace. The profile morphology suggested that it was likely a Lophelia mound. West of this feature, the east face of the east ridge was a steep escarpment running up from 567 to 412 m at the crest. The west ridge crested at 321 m. Total distance from the deep mound to the west ridge was 5.4 km. An E–W echosounder profile at site BU1b, the most southerly site on the Miami Terrace, indicated a double-peaked east ridge crest- ing in 521 m, a valley in 549 m, and the west ridge at 322 m. The east face of the west ridge consisted of a 155-m tall escarpment. Both habitat and faunal zonation varied considerably among the sites. In general, the lower slopes below the ridges and the flat pavement on top of the terrace were relatively barren and had low biodiversity. However, the steep escarpments especially near the tops 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) Octocorallia (Paramuricea spp., Placogorgia spp., Isididae); colonial Scleractinia included scattered thickets of 30–60-cm tall L. pertusa (varying from nearly 100% live to 100% dead), M. oculata (40 cm), and E. profunda; Stylasteridae (15–25 cm), and Antipatharia (30–60 cm) (Table 2). Diverse sponge populations (14 taxa total) of Hexactinellida and Demospongiae included Heterotella spp., Spongosorites spp., Geodia spp., Vetulina spp., Leiodermatium spp., Petrosia spp., Raspailiidae, Astro- phorida, Pachastrellidae, and Corallistidae (Figs. 4, 5). Motile invertebrates included the echinoids Asteroporpa spp. and Stylocidaris spp., the decapod crustaceans C. fenneri and Galatheidae, and other unidentified Echinoidea, Mollusca, and Actini- aria. In total, 19 fish taxa were identified, predominantly shortnose greeneyeChlo- ( rophthalmus agassizi), conger eel (Conger oceanicus), red dory (Cyttopsis rosea), blackbelly rosefish (H. dactylopterus), codling (L. melanurum), dogfish (Mustelus spp.), rattail (Nezumia spp.), dogfish (Triakidae), and dense schools of jacks (Carangi- dae) (Table 3). Schools of ~50–100 wreckfish (P. americanus), ~60–90 cm in length, were observed on several submersible dives in May 2004 and again at the same site in April 2005.

Region I: West Florida Lophelia Lithoherms This region consists of dozens and possibly hundreds of 5–15-m tall lithoherms off the southwest Florida shelf at depths of 500 m, some of which are capped with thickets of live and dead Lophelia. Newton et al. (1987) described the habitat from limited dredge and seismic surveys that extended > 20 km along the shelf slope. In 2003, we conducted a SEABEAM bathymetric survey over a small portion (1.85 × 1.85 km) of the region (Table 1; Figs. 1, 2), and Innovator ROV dives ground- truthed three features: a 36-m tall escarpment and two of the lithoherms. During Innovator ROV dive 6, we examined the 36-m tall escarpment from 412 to 448 m at the eastern edge of the flat terrace that contained the lithoherms. The escarpment REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 367

Table 4. Summary of number of taxa identified for each major faunal group for each study region. Parentheses = standardized number of taxa (taxa/number of dives). (SR = Stetson reefs, SL = Savannah lithoherms, FL = east Florida Lophelia reefs, MT = Miami Terrace escarpment, PT = Pourtalès Terrace, WF = west Florida lithoherms).

Site # of dives Porifera Cnidaria Octocorallia Total sessile taxa Fish SR 7 18 (2.6) 14 (2.0) 6 (0.9) 32 (4.6) 10 (1.4) SL 6 12 (2.0) 10 (1.7) 3 (0.5) 22 (3.7) 10 (1.7) FL 12 18 (1.5) 30 (2.5) 19 (1.6) 48 (4.0) 12 (1.0) MT 11 14 (1.3) 18 (1.6) 8 (0.7) 32 (2.9) 19 (1.7) PT 21 47 (2.2) 22 (1.0) 11 (0.5) 69 (3.3) 30 (1.4) WF 3 6 (2.0) 8 (2.7) 3 (1.0) 14 (4.7) 9 (3.0) Total 60 66 57 32 123 58

was nearly vertical and had very rugged topography with crevices, outcrops, and a series of narrow ledges. Dominant sessile fauna consisted of Antipatharia (30 cm tall), numerous Octocorallia including Isididae (30–40 cm), and sponges, Heterotella spp., Phakellia spp., and Corallistidae. The SEABEAM bathymetry also revealed dozens of lithoherms on a terrace west of the escarpment. Innovator ROV dive 7 was on a 12-m tall and 60-m wide lithoherm (pinnacle 4) in 466 m. Eight other lithoherms were apparent on the ROV’s sonar within a 100-m radius. A transect up the face of the lithoherm revealed a series of terraces on a rugged 45°–70° rock slope that consisted of black rock boulders (1–2 m) and outcrops with 1-m deep crevices. The top ridge was oriented ~NNE–SSW. Thickets of live and dead Lophelia, ~30–60 cm tall and 60–90 cm across, occurred on some terraces but were most abundant on the top ridge. The NE slope appeared to have more live coral than the NW face. Estimated coral cover ranged from <5% to >50% in some areas, with 1%–20% live. The dominant fauna were similar to the escarpment except for Lophelia, which was not observed on the escarpment. Com- mon sessile benthic species included Cnidaria: Antipatharia (Antipathes spp. and Cirrhipathes spp.), L. pertusa, Octocorallia; and Porifera: Heterotella spp. and other hexactinellid vase sponges, and various plate and vase Demospongiae (Pachastrel- lidae, Petrosiidae, Astrophorida) (Table 2, Fig. 5). Common motile invertebrates included Mollusca, Holothuroidea, Crinoidea, and decapod crustaceans (C. fenneri and Galatheidae). Nine species of fish included Anthiinae, shortnose greeneye C.( agassizi), conger eel (C. oceanicus), blackbelly rosefish (H. dactylopterus), codling (L. melanurum), beardfish P( olymixia spp.), and hake (Urophycis spp.) (Table 3). The high number of hard bottom lithoherms revealed by the limited SEABEAM mapping effort and few ROV dives indicate tremendous potential for unexplored coral and fish habitat in this region.

Discussion

Coral Distribution Six regions (B–D, G–I) of deep-water reef habitats were surveyed off southeastern U.S. within the U.S. EEZ from South Carolina to Florida. In addition to these sites, Lophelia reefs and coral have been reported from North Carolina (S.W. Ross, UNCW, pers. comm.; Menzies et al., 1973; Cairns, 1979), in the northeastern Gulf of Mexico on Viosca Knoll and DeSoto Canyon (Schroeder, 2002; Reed et al., 2004; Schroeder et al., 2005), and in the north central Gulf of Mexico in Green Canyon (S.W. Ross, UNCW, 368 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006

pers. comm.; Schroeder et al., 2005). Two regions (E, F) of deep-water reefs are also known in Bahamian waters (Neumann et al., 1977; Mullins et al., 1981; Messing et al., 1990; Reed 2002b), and in 2005, we discovered an isolated 140-m tall Lophelia bioherm at a depth of 780 m west of Great Bahama Bank, south of region F. In our studies, L. pertusa was the dominant colonial scleractinian found at five sites (regions B–D, G, I), but was lacking at the Pourtalès Terrace sites (region H) and the Oculina reefs (region A). Enallopsammia profunda was found in association with Lophelia at all Lophelia reef sites except for the west Florida lithoherm sites (region I). Madrepora oculata was present at the Savannah lithoherms, east Florida Lophelia reefs, and the Miami Terrace escarpment (regions B, C, G). TheLophelia reefs consisted primarily of two habitat types: lithoherms where live coral and coral rubble covered an exposed rock substrate (regions C, D, I), and true Lophelia bioherms where the mounds appeared as coral debris and sediment with little or no exposed bedrock (region B). Paull et al. (2000) estimated that over 40,000 individual lithoherms may cover ~400 km2 on the Blake Plateau and Straits of Florida, perhaps exceeding the areal extent of all the shallow-water reefs of the southeastern U.S. The deep-water reef ecosystems at the Miami and Pourtalès Terraces (regions G, H) were distinctly different from the other Lophelia reef sites. Although some Lophelia colonies were found on the Miami Terrace escarpment, no live colonies were encoun- tered at the Pourtalès Terrace sites (Reed et al., 2005a). The dominant corals at both of these sites were various species of stylasterids with some isidid bamboo octocorals and antipatharian black corals. Various species of isidids were also common at all sites except the Oculina reefs (region A), and black corals were present at all sites.

Deep-water Coral Reef Communities Benthic Communities.—Deep-water coral reefs support rich communities of associated invertebrates. No detailed compilation of benthic fauna has been made previously at these Lophelia reef sites. Historical (Pourtalès, 1868, 1871; Agassiz, 1888) and more recent (Halpern, 1970; Holthuis, 1971, 1974; Cairns, 1979) dredge and trawl surveys of the Blake Plateau and Straits of Florida can only approximate the location of samples or their proximity to deep-water reefs, and none was likely made on steep rock escarpments or under rock ledges. The more recent use of submersibles has allowed for direct observations of the benthic community and the micro-habitat associations of individual species (e.g., Messing, 1984). A total of 142 taxa of benthic invertebrates was identified from these six deep-water reefs (regions B–D, G–I) off the southeastern U.S. Note that some have been identified to species, while others were distinguishable only at generic or familial levels. Although these collections were not quantitative, we targeted the dominant, large (>5cm), sessile organisms, and the numbers of taxa identified were representative of the dominant species at each site. The dominant macrofauna included 66 taxa of Porifera and 57 taxa of Cnidaria. Three taxa were common to all sites: the fan sponge Phakellia spp., Stylasteridae, and the decapod C. fenneri. The octocorals Plumarella pourtalesii and the bamboo coral K. flexibilis were found at all sites except the west Florida lithoherm site. Qualitative differences in species richness were apparent among the sites during the dives, however, direct comparison is difficult since sampling effort differed among the various sites (from 15 dives on the Pourtalès Terrace to three dives on the west Florida lithoherms). Based on number of taxa identified, the Pourtalès Terrace appeared the most species-rich, followed by the east Florida Lophelia reefs, Miami Ter- REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 369

race escarpment, Stetson reefs, and the least was west Florida lithoherm sites (69, 48, 32, 32, and 14 benthic taxa, respectively). However, when numbers were standardized by sampling effort, the west Florida and Stetson reef sites showed greatest faunal richness (4.7 and 4.6, respectively). Of the Lophelia reef sites (regions B–D, G, I), the east Florida Lophelia sites (region B) had the greatest number of cnidarian taxa identified (30), but when these numbers were standardized the west Florida region appeared rich- est (2.7 taxa). In addition, 19 taxa were unique to region B and six were unique to the Stetson reefs. Although the standardized numbers of taxa may indicate that the west Florida lithoherm region had greatest species richness, direct observations during the dives revealed fewer species and abundance than the other deep-reef sites, and it appeared that additional dives would not significantly increase the numbers of dominant taxa. These lithoherms appeared relatively barren except for the upper flanks and ridges where the Lophelia thickets were found. Only 14 sessile taxa were collected, including eight Cnidaria and six Porifera. Certainly, quantitative sampling and the addition of sites at each region are necessary to define the true biodiversity of these deep reefs. Although the Pourtalès and Miami Terraces are adjacent to one another, consider- able differences exist in their benthic communities. Three colonial ScleractiniaE ( nal- lopsammia, Lophelia, and Madrepora) were found on the Miami Terrace escarpment, while we found only S. variabilis on Pourtalès Terrace. Although Lophelia and Mad- repora skeletal debris has been reported from dredge samples on Pourtalès Terrace (Gomberg, 1976; Cairns, 1979), these species were not apparent at our dive sites. Instead of scleractinians, stylasterids dominated the Pourtalès Terrace bioherms and formed thick deposits of live colonies and rubble on their peaks (Reed et al., 2005a). Also strik- ing differences were apparent in the octocorals: only P. pourtalesii was common to both sites, whereas the Miami Terrace had seven unique taxa and Pourtalès Terrace had 10. In addition, we found substantially more sponge taxa on the Pourtalès Terrace than on the Miami Terrace escarpment: 47 and 14 (2.2, 1.3 standardized), respectively. The molluscan fauna were also quite different, with pleurotomariid slit shellsE ( ntem- notrochus adansonianus, Perotrochus amabilis, and Perotrochus midas) and venomous cone snails (Conus villepini) found exclusively on the Pourtalès Terrace. Two types of azooxanthellate corals form deep-water reefs along the Florida coast: O. varicosa and L. pertusa (Reed, 2002a,b). However, the deep-water Oculina reefs are unique to Florida, whereas Lophelia reefs are cosmopolitan. Although these reefs occur at quite disparate depths, 70–100 and 500–800 m, respectively, they are strikingly similar in gross morphology and mound structure due to the similarity of their archi- tectural scleractinians. Both form topographic high-relief features from presumed layers of coral debris and sediment and are capped with living coral thickets. Both occur in regions of strong currents (Florida Current, Gulf Stream), and nutrient-rich, cold- water upwelling exposes the Oculina reefs to periodic low temperatures of 7.4–10 °C, similar to the mean temperatures of the Lophelia reefs in this region. However, the associated fauna are noticeably different between them. For example, 38 taxa of Po- rifera and 41 Cnidaria were identified from theLophelia sites (regions B–D, G–I), but no massive sponges or gorgonians are common to the Oculina bioherms. Live coral coverage is generally low on the majority of both Lophelia and Oculina reefs in this region (1%–10%); however, cover varies from nearly 100% living coral on some reefs to extensive areas of 100% dead coral rubble on others. In previous faunal surveys of the Oculina reefs, collections of coral colonies by lock-out divers from JSL submersibles permitted analyses of the invertebrate community living within the coral. Quantitative 370 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006

collections of 42 Oculina colonies resulted in 20,000 invertebrate specimens including 230 species of mollusks, 50 species of decapods, 47 species of amphipods, 21 species of echinoderms, 15 species of pycnogonids, and numerous unidentified polychaetes and other taxa (Miller and Pawson, 1979; Reed et al., 1982; Pawson and Miller, 1983; Hen- dler and Miller, 1984; Reed and Hoskin, 1987; Reed and Mikkelsen, 1987; Child, 1998). In similar studies of infauna associated with Lophelia in the northeastern Atlantic, Rogers (1999) reported 886 species of coral-associated invertebrates. Quantitative analyses of live and dead colonies of Lophelia off Scotland resulted in 298 species dominated by polychaetes (67 species), bryozoans (45 species), mollusks (31 species), sponges (29 taxa), and crustaceans (15 species) (Jensen and Frederiksen, 1992). In contrast, no quantitative studies have been completed on the infauna associated with Lophelia coral on the western Atlantic reefs. Fish Communities.—In total, 58 taxa of fish were identified from these six deep- water reef sites (regions B–D, G–I), excluding the Oculina reefs. These included 30 species from the Pourtalès Terrace sites, 19 from the Miami Terrace escarpment, 12 from the east Florida Lophelia pinnacles, and ten each from the Savannah lithoherms and Stetson Lophelia reefs. Two species had the greatest occurrence and were found at five sites (blackbelly rose fish, Helicolenus dactylopterus; codling, Laemo- nema melanurum), two species were common to four sites (skate, Raja spp.; rattails, Nezumia spp.), and one species occurred at three sites (shortnose greeneye, C. agassizi). Two species were found solely on the Stetson reef sites, six were unique to the Savannah lithoherm sites, three on east Florida Lophelia sites, seven on the Miami Terrace escarpment, 13 on the Pourtalès Terrace, and two on the west Florida lithoherms. In comparison, the deep-water Oculina reefs appear more species rich (73 species total), but this may reflect a combination of factors including that they occur in substantially shallower and warmer water than Lophelia, they are affected by both coastal water and the Florida Current, species include both shallow-water and deep- shelf taxa, and the Oculina inventory also includes cryptic species collected by lock- out diving from the JSL submersibles (Reed and Gilmore, 1982; Reed et al., 2005b). Nine species were found in common with both the Oculina reefs and the deep-water bioherms on the Pourtalès Terrace, including: Warsaw grouper (Epinephelus nigritus), snowy grouper (Epinephelus niveatus), red barbier (Hemanthias vivanus), ocean sunfish (Mola mola), apricot bass (Plectranthias garrupellus), bank butterflyfish (Prognathodes aya), roughtongue bass (Pronotogrammus matinicensis), greater amberjack (Seriola dumerili), and almaco jack (Seriola rivoliana). However, no species from the Oculina reefs were found on any of the deep-water Lophelia reefs (regions B–D, G, I). Demersal fish communities associated with deep-water habitats in the western At- lantic are poorly known. High-relief, deep-water reefs are not easily trawled and are better sampled by direct observation via submersibles or ROVs (Sedberry and Van Dolah, 1984; Parker and Ross, 1986; Weaver and Sedberry, 2001). Much of the work has concentrated on the Charleston Bump region of the Blake Plateau off Georgia and South Carolina (Sedberry, 2001), which includes our study sites (regions C, D). A combination of factors including upwelling influences of the Gulf Stream, and rugged bottom topography provide essential habitat for various species of commercially important fish including wreck fishP ( . americanus) of which 4.2 million pounds were harvested in 1989 (Sedberry et al., 2001; Vaughan et al., 2001). Using the NR-1 sub- REED ET AL.: FAUNA OF DEEP-WATER LOPHELIA REEFS OFF SE USA 371

marine, Weaver and Sedberry (2001) observed that 15 species of fish were commonly associated with hard-bottom habitats on the Charleston Bump. Of these, 11 taxa were observed at our study sites. Our sites at regions C and D also resulted in several taxa not listed by Weaver and Sedberry (2001) including the gaper (Chaunax spp.), roughies (Hoplostethus mediterranus), hagfish (M. glutinosa ), hake (Phycis spp.), skate (Raja spp.), and cutthroat eel (Synaphobranchidae). North of region B lies the Agassiz Hills (650–750 m), which are hard-bottom reefs dominated by solitary corals (Bathypsammia spp. and Flabellum spp.) rather than Lophelia (George, 2002). These also have high fish biodiversity due in part to episodic upwelling. Bottom trawls at the Agassiz Hills collected 24 species of demersal fish including commercially important rattail fish Coryphenoides( armatus), deep sea eel (Synaphobranchus kaupi), and wreck fish P( . americanus). Of these, eight taxa overlap with our Lophelia reef sites, including hagfish (M. glutinosa), lantern shark (Etmopterus spp.), shortnose greeneye (C. agassizi), shortbeard codling (Laemonena barbatulum), hake (Urophy- cis spp.), rattails (Nezumia spp.), and wreckfish P( . americanus). George (2002) com- pared the Agassiz Hills with a second site off north Florida (650–800 m), which is within our region C, but only four fish species were collected, whereas we found 12 species on the east Florida Lophelia reefs.

Management of Deep-Sea Ecosystems Region A, the deep-water Oculina reefs, has been designated a habitat area of particular concern (HAPC) for the protection of the coral habitat (Reed and Gilmore, 1982; Reed, 2002b; Reed et al., 2005b). A portion also has been designated a marine protected area (MPA) for management of the snapper-grouper complex. Approxi- mately one-third of the reef system (315 km2; 92 nm2) was first designated as a HAPC in 1984, and was then expanded northward to encompass 1029 km2 (300 nm2) in 2000. Even so, extensive areas of the Oculina reefs have been severely impacted by both legal and illegal bottom trawling since 1984, especially in the later protected area (Koenig et al., 2005; Reed et al., 2005b). Some areas in the northern section of the MPA that were documented as thriving reefs by photo transects in the 1970s had been found to be reduced to 100% rubble during submersible dives in 2001 (Reed et al., 2005b). However, some of the reefs in the southern portion that had been protected since 1984 are still thriving with nearly 100% live coral cover. In comparison, the Lophelia reef sites vary greatly in live coral coverage, from nearly 100% live on reefs at Viosca Knoll and off North Carolina to other sites that are < 10% live. So far we have no evidence that commercial bottom trawling has occurred on the Lophelia reefs in this region of the western Atlantic, and so it is still speculative as to whether the cause of the high percentage of dead coral could be due to natural senescence of the reefs, paleoclimatic factors, coral pathogens, or other unknown factors. The six deep-water reefs outlined here (regions B–D, G–I) are each unique and their potential for fisheries and biopharmaceutical resources remain relatively unknown. Protection is needed to prevent long-term (perhaps permanent) damage, such as what has occurred on the Oculina reefs off Florida and on theLophelia reefs in the northeastern Atlantic, both destroyed in part by trawling (Rogers, 1999; Fosså et al., 2002; Koenig et al., 2005; Reed et al., 2005b). Because trawlers are banned from the Oculina HAPC, concern exists that trawlers may move to deeper habitats in search of valuable commercial fisheries, such as royal red shrimp or benthic fin- fish. Evidence of aggregations of wreckfishP ( . americanus), populations of blackbelly 372 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 2, 2006 rosefish H( . dactylopterus), and other commercially important species could threat- en the longevity of these fragile habitats unless bottom trawling in these regions is prohibited or strictly regulated and monitored. Activities involving pipelines or oil/ gas production could also negatively impact these reefs. Although these habitats are not currently designated as MPAs or HAPCs, they are under consideration for such status by the South Atlantic Fishery Management Council and NOAA Fisheries. The surveys summarized in this report are only preliminary and point to the need for additional geological, biological, and ecological research. As a first step, detailed mapping and habitat characterization studies would provide data for designation of potential MPA and HAPC boundaries and future research needs.

Acknowledgments

Numerous individuals have contributed to this research over many years. R. Avent initi- ated the submersible studies of Florida’s deep-water reefs in the 1970s and C. Hoskin provided years of enthusiastic collaboration and leadership. We especially thank Harbor Branch Oceanographic Institution (HBOI) and the Division of Biomedical Marine Research for fund- ing submersible and ship time for these studies. NOAA’s Office of Ocean Exploration funded our expeditions for biomedical research on these reefs in 2002 and 2003. The State of Florida funded the Center of Excellence in Biomedical and Marine Biotechnology (HBOI and Florida Atlantic University), which provided ship and submersible time in 2004 and 2005. The following individuals assisted with taxonomic identifications: Porifera- M. Kelly (National In- stitute of Water and Atmospheric Research, New Zealand), K. Ruetzler (National Museum of Natural History); Cnidaria- S. Cairns (National Museum of Natural History); Echinoderms- C. Messing (Nova Southeastern Univ.), J. Miller (HBOI), D. Pawson (National Museum of Natural History); Mollusca- P. Mikkelsen (American Museum of Natural History), A. Olenik (Florida Atlantic University), M. Vecchione (National Museum of Natural History); Fish- R.G. Gilmore (ECOS), C.R. Robins (formerly RSMAS), J. McEachran (Texas A&M University). We especially thank R.G. Gilmore for his contributions to the knowledge about fish communties on the Oculina reefs and providing the Oculina fish species list. The various crews of HBOI research vessels, and Johnson-Sea-Link and Clelia submersibles are gratefully thanked for their support, without which this research would not have been possible. S. Reed (Smithson- ian Marine Station) and C. Messing are thanked for their comments and reviews on drafts of the manuscript. This is Contribution Number 1610 from Harbor Branch Oceanographic Institution and P200604 from the Center of Excellence in Biomedical and Marine Biotech- nology.

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Date Submitted: 22 February, 2005. Date Accepted: 11 October, 2005.

Addresses: (J.K.R., S.A.P.) Harbor Branch Oceanographic Institution (HBOI), 5600 U.S. 1, North, Fort Pierce, Florida 34946. (D.W.) Flower Garden Banks National Marine Sanctuary NOS/NOAA, 1200 Briarcrest Drive, Suite 4000, Bryan, Texas 77802. Corresponding Au- thor: (J.K.R.) E-mail: .