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. Washington, DC20013-7012 MRC-153 Smithsonian Inst., P.O. Box37012, NHB, WC-57, Systematics Laboratory 2 Watershed Studies,StPetersburg,FL US GeologicalSurvey, CenterforCoastal & Personnel Act) to: *Currently assigned(throughIntergovernmental Wilmington, NC28409 5600 MarvinMossLn. 1 NationalMarineFisheries ServiceNational UNC-Wilmington,CenterforMarineScience Oculina varicosa Oculina INTRODUCTION Lophelia CAPE HATTERAS TO SOUTHEASTERNFLORIDA Oculina .

pertusa ,

STATE OFDEEP CORAL ECOSYSTEMS and the second includes second the and ak, eeecs on references banks, IN THEU.S.SOUTHEASTREGION: pu ohr corals other (plus 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 this unique deep-reef habitat. Yet, it is 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 Although forms 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 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 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 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 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 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 . collected corals on the Blake Plateau Blake the on corals collected and , (Agassiz 1888). These collections (= (= Balanophyllia Caryophyllia Dendrophyllia 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 ) Tethocyathus squiresi )

profunda Lophelia pertusa , . , ,

ugse 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 and of occurrence “Stetson Banks.” Two of these dives confirmed the DSRV the using dives submersible manned five 1967, 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 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 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 cruise, S.W. Ross, chief scientist), but no hard no but scientist), chief S.W.Ross, cruise, Eastward Lophelia Eastward Alvin ee ae n n ra et f the of west area an in made were in the region (Milliman et al. 1967). al. et (Milliman region the in s et sudr LR McCloskey (L.R. sounder depth ’s tto ae ws rwe and trawled was area station Enallopsammia Eastward Lophelia Lophelia Alvin cruises to this area off area this to cruises NR-1 p ocre of the off occurred sp. dives. off Cape Lookout. Cape off nuclear research nuclear (= Eastward Dendrophyllia Delaware (E-25- )

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 1970;1980), 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 rdd msl drn te litcn) y the 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 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. 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, Gulf 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 Wilmington. NOAA UnderseaResearchCenteratUNC- the Florida Figure 6.2. Oculina 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 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 (<200 shelf varicosa Oculina outer the U.S. on coral southeastern structure-forming dominant The Scleractinia) a. StonyCorals

ioy re oa) lhuh it Although coral). tree (ivory Oculina (Class Anthozoa, Order 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 Fear Cape Lookout . Depthsounderprofilesand selectedbottomviewsofthethreeNorthCarolina Lophelia 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 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 Madrepora oculata is still dominant. For example, the colonial corals lished data. yet beenidentified.Photocredit:Rossetal.unpub south ofNorthCarolina.Othercorals(C)havenot sp.) coralscommonlyfoundontheBlakePlateau toisis possibly Figure 6.5. C A spp.,possibly and Leiopathes glaberrima Examplesofblack(A,B; ahpama fallosocialis Bathypsammia K. ornata and Enallopsammia profunda ) andgold(E; ), bamboo(D; STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Thecopsammia Leiopathes Gerardia Lophelia carpet Kera 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 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 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 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 the ( 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) d. Gorgonians huel bipinnata Thourella (Class Anthozoa, Order hyooga squamata Chrysogorgia (Class Anthozoa, Order 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 , , lhuh o fud n ra abundances, great in found the off occur commonly corals) (lace stylasterids not Although Anthoathecatae) g. Stylasterids ( species sertum 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 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 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 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. increase southwardfromtheCarolinas. stylasterids of diversity and Abundance rubble. often attached to dead scleractinian corals or coral observed Individuals 6.1). (Appendix region the from reported been have genera four representing species Seven U.S. southeastern ) isknownintheregion(Appendix6.1). CORAL COMMUNITIES SPECIES ASSOCIATIONS WITHDEEP banks is typical of the southeastern U.S. Oculina (ClassHydrozoa,Order (Class Anthozoa, Order 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 Photographs credit:S.W. Ross. Figure 6.6 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 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) abundant and al. et 2006). Reed diverse and 6.3 (Table fauna the invertebrate is corals the from themselves) (aside habitats coral these of 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 et al.unpublisheddata,2=Reed2006,3=Henryinreview. southeastern U.S. deep coralhabitats.Coralsarelistedseparatelyin Appendix 6.1. References are1=Nizinski Table 6.3. Phylum Arthopoda Phylum Mollusca Phylum Porifera(Sponges) Subphylum Crustacea Class Malacostraca Class Gastropoda Class Cephalopoda Class Hexactinellida(glasssponges) Class Demospongiae Order Decapoda Coralliophila Octopus, multiplespecies Squids, Aphrocallistes beatrix multiple species multiple species Preliminarylistofdominantbenthicmegainvertebratesobservedorcollectedonnear Other taxa Infraorder Brachyura Infraorder Anomura and theirrelatives) Superfamily Paguroidea(hermitcrabs Shrimps, multiplespecies Family Portunidae Family Geryonidae Family Pisidae Family Galatheidae(squatlobster) Family Chirostylidae(squatlobster) multiple species Ilex Chaceon fenneri crab) Rochinia crassa Munidopsis Munida Uroptychus Gastroptychus salvadori Eumunida picta swimming crab) Bathynectes longispina crab) sp. (?)sp. 1,2 1 1

1,2 1,2 spp. including 1 spp. 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 Crinoidea(crinoids) Class Polychaeta(polychaetes) Class Anthozoa Class Hydrozoa(Hydroids) Class Echinoidea(seaurchins) Class Ophiuroidea(brittlestars) Class Asteroidea (seastars) Order Zoanthidea(zoanthids) Order Actinaria (anemones) Order Cidaroida Order Echinothurioida Order Echinoida Order Brisingida(brisingidseastar) multiple speciesincluding multiple species(≥37species) bidentata multiple species multiple species multiple species multiple species rugosa multiple speciesincluding Family Cidaridae Family Echinothuriidae Family Echinidae Family Brisingidae Stylocidaris Cidaris rugosa Hygrosoma E. tylodes Echinus gracilis Novodinia antillensis (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 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 the impact to range depth a within area ( lionfish the species, invasive one only date To impact tocorals(Guinotteetal.2006). 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 (except deep such region impacts southeast the to of corals impacts sedimentation documenting Table 6.4 Pterois volitans Bottom-set Longline Bottom-set gillnet Mid-water Trawl Traps orPots Bottom Trawl Gear Type Dredge . PotentialfishinggearimpactstodeepwatercoralsinthesoutheasternUnitedStates. ), 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. 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. 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 Low Low Low Low Low Low MANAGEMENT OFFISHERY RESOURCES AND HABITATS Oculina www.safmc.ne ak. ee seis are species Fewer banks. Overall Ratingof Gear Impact Medium High Low Low Low Low t . aaeet is Management ). 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   Madrepora 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 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 Atkinson LP, Pietrafesa LJ, Hofmann EE (1982) EE Hofmann LJ, LP,Pietrafesa Atkinson Arendt MD, Barans CA, Sedberry GR, Van Dolah United the of cruises Three (1888) A Agassiz X. 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 a, ot Crln. ora o 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 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 REFERENCES

255 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Bane JM Jr, Attkinson LP, Brooks DA (2001) Buhl-Mortensen L, Mortensen PB (2004) Gulf Stream physical oceanography at the Symbiosis in deep-water corals. Symbiosis Charleston Bump: deflection, bimodality, 37:33-61 meanders, and upwelling. In: Sedberry GR (ed) Island in the Stream: oceanography Cairns SD (1979) The deep-water scleractinia and fisheries of the Charleston Bump. Amer of the Caribbean Sea and adjacent waters. Fish Soc Symposium 25, Bethesda, MD, p Studies on the fauna of Curacao and other 25-36 Caribbean Islands 57(180), 341 pp

Bayer FM (2001) New species of Calyptrophora Cairns SD (1981) Marine flora and fauna of the (Coelenterata: Octocorallia: Primnoidae) Northeastern United States, Scleractinia. from the western part of the Atlantic Ocean. NOAA Technical Report NMFS Circular 438, Proceedings of the Biological Society of 14 pp Washington 114:367-380 Cairns SD (1986) A revision of the Northwest SOUTHEAST Blake JA, Hecker B, Grassle JF, Brown B, Wade Atlantic Stylasteridae (Coelenterata: M, Boehm PD, Baptiste E, Hilbig B, Maciolek Hydrozoa). Smithsonian Contributions to N, Petrecca R, Ruff RE, Starczak V, Watling Zoology 418, 131 pp L (1987) Study of Biological Processes Cairns SD (1997) A generic revision and on the U.S. South Atlantic Slope and phylogenetic analysis of the Turbinoliidae Rise: Phase 2. US Department of Interior, (Cnidaria: Scleractinia). Smithsonian Minerals Management Service, Atl OCS Contributions to Zoology 591, 55 pp Region, Environmental Assessment Section. Herndon, VA, 414 pp + appendices Cairns SD (2000) A revision of the shallow-water azooxanthellate Scleractinia of the western Borowski WS, Paull CK, Ussler W III (1997) Carbon Atlantic. Studies on the Natural History of cycling within the upper methanogenic zone the Caribbean Region 75, 240 pp of continental rise sediments: an example from the methane-rich sediments overlying Cairns SD (2001a) A generic revision and the Blake Ridge gas hydrate deposits. phylogenetic analysis of the Dendrophylliidae Marine Chemistry 57:299-311 (Cnidaria: Scleractinia). Smithsonian Contributions to Zoology 615, 75 pp Brooke S, Young CM (2003) Reproductive ecology of a deep-water scleractinian coral, Cairns SD (2001b) Studies on western Atlantic Oculina varicosa, from the southeast Florida Octocorallia (Coelenterata: Anthozoa). Part shelf. Continental Shelf Research 23:847- 1: The genus Chrysogorgia Duchassaing 858 & Michelotti, 1864. Proceedings of the Biological Society of Washington 114:746- Brooke S, Young CM (2005) Embryogenesis and 787 larval biology of the ahermatypic scleractinian Oculina varicosa. Marine Biology 146:665- Cairns SD (2006) Studies on western Atlantic 675 Octocorallia (Coelenterata: Anthozoa). Part 6: The genera Primnoella Gray, 1858; Brooks RA, Nizinski MS, Ross SW, Sulak KJ Thouarella Gray, 1870; Dasystenella (2007) Frequency of sublethal injury in a Versluys, 1906. Proceedings of the Biological deepwater ophiuroid, Ophiacantha bidentata, Society of Washington 119:161-194 an important component of western Atlantic Lophelia reef communities. Marine Biology Cairns SD, Bayer FM (2002) Studies on 152:307-314 western Atlantic Octocorallia (Coelenterata: Anthozoa). Part 2. The genus Callogorgia Brundage WL (1972) Patterns of manganese Gray, 1858. Proceedings of the Biological pavement distribution on the Blake Plateau. Society of Washington 115:840-867 In: Horn DR (ed) Ferromanganese deposits on the ocean floor. National Science Cairns SD, Bayer FM (2003) Studies on Foundation. Washington, DC, pp 221-250 western Atlantic Octocorallia (Coelenterata:

256 oea F, ong C Eln -, Grimes A-M, Eklund CC, Koenig FC, Coleman T,Morato TJ, Pitcher R, Watson WWL, Cheung, GR Sedberry KJ, Sulak SW, Ross JH, Caruso Cairns SD, Hoeksema BW, van der Land J (1999) Biogeographic (2001) RE Chapman SD, Cairns on Studies (2004b) FM Bayer SD, Cairns Studies (2004a) FM Bayer SD, Cairns mrcn ihre Scey Symposium Society 23:233-242 Fisheries American States. United southeastern the of complex temperate reef fishes in the grouper-snapper CB (1999) Management and conservation of Series 333:1-12 Progress Ecology Marine catch. fish global the in vulnerability Intrinsic (2007) D Pauly of Journal Fisheries Biology70:1015-1026 States. United south-eastern off the Lophiiformes) (Pisces: anglerfishes lophiid and chaunacid Deep-water (2007) Research Bulletin459:13-46 Atoll corals. stony extant of List Appendix: Nova Scotia,pp30-57 Ecology Halifax, Centre, Corals. Action sea Deep- on Symposium Int First Proc. (eds) P Doherty M, Butler ELR, Kenchington SE, Gass J, deep-water Hall JHM, Willison In: Scleractinia. Atlantic North the of affinities 487 117:447- Washington of Society Biological Candidella 1870; Gray, genera The 5. Part Anthozoa). (Coelenterata: Octocorallia Atlantic western of Octocorallia Society Washington 117:114-139 Biological the of The Atlantic Proceedings 4. Part genus Anthozoa). western (Coelenterata: on of Washington 116:617-648 1870. Proceedings of the Biological Society 3. Part genus Anthozoa). The Paracalyptrophora Bayer, 1954. Proceedings of the Acanthoprimnoa ioht, 1908. Kinoshita, STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF n gn; and gen.; n. , Narella Plumarella Gray,

Druffel ERM, Griffin S, Witter A, Nelson E, A,Nelson Witter S, Griffin ERM, Druffel Buesseler RA, Belastock LL, King ERM, Druffel WP,Dillon Plateau Blake The P(1988) Popenoe T,Lundalv A, Freiwald M, McCrea MJ, Costello ate J J, ua K, os W Necaise SW, Ross KJ, Sulak Jr, JV Gartner (2002) DM Furevik PB, Mortensen JH, Fossa new A press) (in AM Quattrini B, Fernholm (1978) RY George JH, Hudson C, Emiliani of Atlas (1991) Staff Scientific 87 EEZ-SCAN et Cosmochimica Acta 54:1493-1500 using coral deep-sea a of rate Growth (1990) KO North of America, p291-327 Society Geological The 1-2. vol Margin, U.S. The Geology of North America, Sheridan In: JACrow (eds) The RE, Continental Atlantic Trough. Carolina and Basin Springer- ecosystems. Verlag BerlinHeidelberg and corals water Cold- (eds.), JM Roberts Freiwald A, in 805 Pages 771- NEAtlantic. the in habitat fish cold-water of JM, Role Roberts (2005) H, D Haas Allen de TCE, Weering van BJ, Brett L, Jonsson slopes. MarineBiology continental Virginia and Carolina North the near-bottom Persistent along animals mesopelagic of aggregations review) (in A-M impacts. Hydrobiologia471:1-12 fishery and distribution waters: Norwegian coral deep-water The western North Atlantic. Copeia associated with deep-sea coral habitat in the Eptatretus) (Myxinidae: hagfish of species 202:627-629 deep-sea coral from Blake Plateau. Science a and Keys Florida the from coral reef a in record growth isotope carbon and Oxygen Miscellaneous InvestigationsSeriesI-2054 Survey Geological US margin. continental Atlantic zone, economic exclusive U.S. the 5036 Geochimica et Cosmochimica Acta 59:5031- Gerardia (1995) J Vogel M, Kashgarian J, Southon 210 Pb and other isotopes. Geochimica : bristlecone pine of the deep-sea? the of pine bristlecone : ohla pertusa Lophelia ohla pertusa Lophelia oa res as reefs coral in

257 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Genin A, Dayton PK, Lonsdale PF, Spiess FN Hovland M, Risk M (2003) Do Norwegian deep- (1986) Corals on seamount peaks provide water coral reefs rely on seeping fluids? evidence of current acceleration over deep- Marine Geology 198:83-96 sea topography. Nature 322:59-61 Jensen A, Frederiksen R (1992) The fauna Genin A, Paull CK, Dillon WP (1992) Anomalous associated with the bank-forming deepwater abundances of deep-sea fauna on a rocky coral Lophelia pertusa (Scleractinaria) on bottom exposed to strong currents. Deep- the Faroe Shelf. Sarsia 77:53-69 Sea Research 1 39:293-302 Koenig CC, Coleman FC, Grimes CB, Fitzhugh George RY (2002) Ben Franklin temperate GR, Scanlon KM, Gledhill CT, Grace M reef and deep sea “Agassiz Coral Hills” (2000) Protection of fish spawning habitat in the Blake Plateau off North Carolina. for the conservation of warm-temperate reef- Hydrobiologia 471:71-81 fish fisheries of shelf-edge reefs of Florida.

SOUTHEAST Bulletin of Marine Science 66:593-616 Gilmore RG, Jones RS (1992) Color variation and associated behavior in the epinepheline Koenig CC, Shepard AN, Reed JK, Coleman FC, groupers, Mycteroperca microlepis (Goode Brooke SD, Brusher J, Scanlon KM (2005) and Bean) and M. phenax Jordan and Swain. Habitat and fish populations in the deep- Bulletin of Marine Science 51:83-103 sea Oculina coral ecosystem of the Western Atlantic. In: Barnes PW, JP Thomas (eds) Griffin S, Druffel ERM (1989) Sources of carbon Benthic Habitats and the Effects of Fishing. to deep-sea corals. Radiocarbon 31:533- American Fisheries Society Symposium 41, 543 Bethesda, MD, p 795-805

Guinotte JM, Orr J, Cairns S, Freiwald A, Morgan Koslow JA (1997) Seamounts and the ecology L, George R (2006) Will human-induced of deep-sea fisheries. American Scientist changes in seawater chemistry alter the 85:168-176 distribution of deep-sea scleractinian corals? Frontiers Ecology and Environment 4:141- Koslow JA, Boehlert GW, Gordon JDM, Haedrich 146 RL, Lorance P, Parin N (2000) Continental slope and deep-sea fisheries: implications Gulf of Mexico Fishery Management Council and for a fragile ecosystem. ICES Journal of South Atlantic Fishery Management Council Marine Science 57:548-557 (1982) Fishery Management Plan Final Environmental Impact Statement for Coral Le Goff-Vitry MC, Pybus OG, Rogers AD (2004) and Coral Reefs. Charleston, SC Genetic structure of the deep-sea coral Lophelia pertusa in the northeast Atlantic Hain S, Corcoran E (eds) (2004) 3. The status of revealed by microsatellites and internal the cold-water coral reefs of the world. Pages transcribed spacer sequences. Molecular 115-135. In: Wilkinson C (ed) Status of coral Ecology 13:537-549 reefs of the world: 2004. vol 1. Australian Inst Mar Sci Perth, Western Australia Lee TN, Yoder JA, Atkinson LP (1991) Gulf Stream frontal eddy influence on productivity of the Henry L-A, Nizinski MS, Ross SW (in press) southeastern U.S. continental shelf. Journal Diversity, distribution, and biogeography of of Geophysical Research 96:22191-22205 hydroid assemblages collected from deep- water coral habitats off the southeastern Macintyre IG, Milliman JD (1970) Physiographic United States. Deep-sea Research I features on the outer shelf and upper slope, Atlantic continental margin, southeastern Hovland M, Mortensen PB, Brattegard T, Strass United States. Geological Society of America P, Rokoengen K (1998) Ahermatypic coral Bulletin 81:2577-2598 banks off mid-Norway: evidence for a link with seepage of light hydrocarbons. Palaios 13:89-200

258 Mikkelsen N, Erlenkeuser H, Killingley JS, Berger (1990) JC Lang AC, Neumann CG, Messing (1973) GT Rowe RY, George RJ, Menzies Ross JK, Loefer DM, Wyanski HS, Meister Summary: (2003) KA Puglise JJ, McDonough New A press) (in SW Ross JE, McCosker and Summer (1986) O Pashuk TD, Mathews CL, Jacobs DSM, Billett BJ, Bett DG, Masson Structure (1970) JI Ewing GM, Bryan RG, Markl Geomorphology (1970) R Hurley RJ, Malloy Boreas 11:163-171 in isotopes stable and radiocarbon corals: Norwegian (1982) WH Straits ofFlorida.Palaios5:15-33 northeastern the in hardgrounds associated and lithoherms deep-water of Biozonation York, USA New Sons, and the Wiley John Oceans. World of Ecology and Environment Abyssal SE Nat4:193-206 along the Atlantic coast of the United States. of evidence for the invasion and establishment Further (2005) KJ Sulak AM, Quattrini SW, Tech MemoNMFS-SPO-60,51pp Galway, Ireland, January 16-17, 2003. NOAA Gulf of Mexico and the North Atlantic Ocean. International the for workshop collaboration and planning workshop. corals Deep-sea from NorthCarolina.Copeia Ophichthus Deepwater Species of the Snake Eel Genus Coastal of Journal Research 2:311-336 (1973-1979). Bight winter hydrography of the U.S. South Atlantic Marine Geology194:159-180 Trough,Rockall Northeast northern Atlantic. the in mounds origin topped coral deep-water, The of (2003) RB Wynn AJ, Wheeler Geophysical Research75:4539-4555 of the Blake-Bahama outer ridge. Journal of Bulletin America of 81:1947-1972 Society Florida. Geological of Straits structure: geologic and Pterois volitans Agilfre: Ophichthidae) (Anguilliformes: (Teleostei: Scorpaenidae) ohla pertusa Lophelia STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION U.S. SOUTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF . Morgan LE, Tsao C-F, Guinotte JM (2006) Status and Oxygen (1998) HT Rapp PB, Mortensen Milliman JD, Manheim FT, Pratt RM, Zarudzki EFK area protected Marine (2001) CA Miller itaea J 18) hlbek circulation, Shelfbreak (1983) LJ Pietrafesa Ussler BA, Ende am AC, Neumann CK, Paull Sedberry AM, Quattrini SW, Ross ML, Partyka (1977) G Keller JW, Kofoed AC, Neumann Submersible (1970) MM Ball AC, Neumann Sarsia 83:433-446 rates. extension linear implications of determination for Scleractinia): (Anthozoa, (L) of skeletons in patterns line growth to related ratios isotope carbon Technical ReportRefNo.67-80 Institute Oceanographic Hole Woods 1967. off the southeastern United States July 2-13, (1967) margin continental the on dives Alvin Nova ScotiaMuseum.Halifax, on Deep-Sea Corals. Ecology Action Centre. Symposium International First Proc. (eds). P Doherty M, Butler ELR, Kenchington SE, Gass J, Hall JHM, Willison In: 145-155. p conservation. coral deep-sea for framework 250 33:233- Publication Special and Mineralogists Paleontologists Economic margins. of continental Society on interface critical In: perspectives. coast west shelfbreak: The and east oceanography: physical and fronts 166:83-101 Geology Marine slope. Florida-Hatteras the on Lithoherms (2000) NM Rodriguez III, W Silver Spring,MD,176pp NOAAcorals. deep-sea Tech1. OER Memo effort to characterize areas of habitat-forming Corals (SEADESC) Initiative: a collaborative press) Southeastern United (in States Deep S Sea Gottfried J, Potter TW, Birdsong GR, 5:4–10 Lithoherms in the Straits of Florida. Geology America Bulletin81:2861-2874 of Society Geological currents. bottom and observations in the Straits of Florida: geology Conservation Marine Biology Institute,Bellevue,WA, 63pp Management. Conservation and with their Waters, for US in Recommendations Corals Sea Deep of ohla pertusa Lophelia

259 SOUTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Pinet PR, Popenoe P, Otter ML, McCarthy SM Reed JK (1981) In situ growth rates of the (1981) An assessment of potential geologic scleractinian coral Oculina varicosa hazards of the northern and central Blake occurring with zooxanthellae on 6-m reefs Plateau. In: Popenoe P (ed) Environmental and without on 80-m banks. Proceedings of geologic studies on the southeastern Atlantic the 4th International Coral Reef Symposium outer continental shelf. USGS Open File 2:201-206 Rept 81-582-A, pp 8-1-8-48 Reed JK (1983) Nearshore and shelf-edge Popenoe P (1994) Bottom character map of the Oculina coral reefs: the effects of upwelling northern Blake Plateau. USGS Open File on coral growth and on the associated faunal Rept 93-724 communities. NOAA Symposium Series Undersea Research 1:119-124 Popenoe P, Manheim FT (2001) Origin and history of the Charleston Bump-geological Reed JK (2002a) Comparison of deep-water

SOUTHEAST formations, currents, bottom conditions, and coral reefs and lithoherms off southeastern their relationship to wreckfish habitats on the U.S.A. Hydrobiologia 471:57-69 Blake Plateau. In: Sedberry GR (ed) Island in the Stream: oceanography and fisheries Reed JK (2002b) Deep-water Oculina coral of the Charleston Bump. American Fisheries reefs of Florida: biology, impacts, and Society Symposium 25, Bethesda, MD, p management. Hydrobiologia 471:43-55 43-93 Reed JK (2004) Deep-water coral reefs of Florida, Pratt RM (1968) Atlantic continental shelf and Georgia and South Carolina: a summary slope of the United States - physiography of the distribution, habitat, and associated and sediments of the deep-sea basin. U S fauna. Unpublished Rept to South Atlantic Geological Survey Professional Paper529- Fishery Management Council, Charleston, B, 44 pp SC, 71pp

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 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 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 (2006) SA Pomponi DC, Weaver JK, Reed 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 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 water 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 Lnau 15) n ohr deep- other and 1758) (Linnaeus 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 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 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 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 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, W,Knick 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

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

Wenner EL, Barans CA (2001) Benthic habitats and associated fauna of the upper- and middle-continental slope near the Charleston Bump. pp 161-178 In: Sedberry GR (ed) Island in the Stream: oceanography and fisheries of the Charleston Bump. American Fisheries Society Symposium 25, Bethesda, MD

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; 220-1383 403-1748 300-1646 Depth Range commonly 500- SOUTHEAST Distribution Straits; eastern Nova Scotia - FL Antilles Gulf of Mexico; Lesser - Suriname GA Straits of FL; northern Gulf Arrowsmith Bank, of Mexico; Yucatan Arrowsmith Bank, NC, GA; Yucatan - Straits of Florida MA 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 1848) Paracyathus pulchellus Premocyathus cornuformis Solenosmilia variabilis Stephanocyathus coronatus Stephanocyathus diadema Stephanocyathus laevifundus Stephanocyathus paliferus cylindraceus Tethocyathus recurvatus Tethocyathus variabilis Tethocyathus rawsonii Trochocyathus Balanophyllia cyathoides Balanophyllia floridana Bathypsammia fallosocialis Bathypsammia tintinnabulum Cladopsammia manuelensis Eguchipsammia gaditana Enallopsammia profunda Enallopsammia rostrata 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 41 MR 25 MR 63 MR 19 MR 44 MR 42 MR 27 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 24-329 29-861 13-105 24-134 175-586 320-3186 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 (Nova Scotia, Newfoundland); MA; DE; GA; FL; Bahamas Nova Scotia, Canada (off NC Newfoundland); MA; VA; FL; Bahamas; Colombia; Tobago; Trinidad; Venezuela; Suriname; Dominican Republic; St. Lucia Nova Scotia, Canada (off Newfoundland); ME; MA; SC; GA; FL SC; Bahamas; Dominican Republic; Martinique NC; SC; GA; FL Havana, (off NC; Straits of FL Cuba); Bahamas FL, SC; FL; Gulf of Mexico (off LA) FL Nova Scotia, Canada (off FL Newfoundland); MA; DE; VA; SC; GA; FL; Bahamas; Straits of Keys; Havana, Cuba); FL (off FL Keys) FL Gulf of Mexico (off TX) FL; Gulf of Mexico (FL; GA; FL; Bahamas; Straits of FL Gulf of Mexico Key West); (off Antilles; Brazil Keys); Lesser (FL Jacksonville, Fl; Caribbean 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, Anthomastus agassizi Anthomastus grandiflorus Bellonella rubistella Clavularia modesta Scleranthelia rugosa fruticulosa Telesto nelleae Telesto sanguinea Telesto rudis Trachythela Gersemia fruticosa Pseudodrifa nigra Siphonogorgia agassizii Chrysogorgia multiflora Chrysogorgia squamata Higher Taxon Subclass Octocorallia Alcyonacea Order Alcyoniidae Family Family Clavulariidae Family Nephtheidae Family Nidaliidae Order Gorgonacea Family Chrysogorgiidae

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 2004b 2004a 2004a 2004b 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, 2002 C & B, 2003 C & B, 2003 C & B, 2003 82-514 593-911 221-858 183-732 229-556 161-792 677-900 374-555 165-706 310-878 348-572 514-2063 738-1473 494-1065 Depth Range SOUTHEAST Distribution Lydonia Canyon; FL; Gulf of Lydonia Keys) FL Mexico (off Antilles Straits of Florida; Lesser central Florida; Bahamas; off Honduras; northern Antilles; off Gulf of Mexico Straits of FL SC; GA; Bahamas New England seamounts; Bermuda; eastern coast FL; Antilles; northern Gulf Bahamas; of Mexico Delray Beach); (off Straits of FL Antilles Bahamas; Lesser Delray Beach); (off Straits of FL Bahamas; Cuba; Puerto Rico; Campeche Bank, Mexico St. (off Bermuda; Straits of FL Lucie Inlet, Palm Beach, Delray Beach;Bahamas); Cuba Cape Canaveral (off Straits of FL - Cuba); Bahamas; Lesser Antilles (off Insular side Straits of FL Palm Beach, north of Little Bahama Bank), Bahamas to Channel Yucatan SC to Cuba off SC to FL off North and South Carolina off 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 Swiftia koreni Callogorgia americana 2002 Bayer, Callogorgia gracilis 1857) Calyptrophora gerdae Calyptrophora trilepis 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