STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION: MAINE TO CAPE HATTERAS

David B. Packer1, Deirdre Boelke2, Vince Guida1, and Leslie-Ann McGee2

I. INTRODUCTION Mortensen and Buhl-Mortensen 2004; Gass and Martin Willison 2005; Mortensen et al. 2005). The U.S. Northeast Shelf Large Marine Ecosystem Many gorgonian corals such as Paragorgia have encompasses 260,000 square km and extends been regularly encountered by fishermen on from the Gulf of Maine and Georges Bank hard substrate such as boulders, gravel, or rocky southward through southern New England waters outcrops (e.g., Breeze et al. 1997; Leverette and NORTHEAST and the Middle Atlantic Bight to Cape Hatteras. It Metaxas 2005). extends from the coast eastward to the edge of the continental shelf and slope and offshore to This chapter summarizes the current knowledge the Gulf Stream (Sherman et al. 1996). Some of of deep corals and deep coral communities for the the specific locations with known occurrences of U.S. Northeast Shelf Large Marine Ecosystem, deep corals include parts of the Gulf of Maine, including the diversity of coral species and their Georges Bank, as well as a number of canyons distribution, associated species, and habitat that bisect the continental shelf and slope. Deep preferences. In addition, a summary of current corals have also recently been collected from the stressors affecting these habitats, and existing New England Seamount chain; the seamounts conservation and management activities are are located off the continental shelf, rising above presented. Gaps in our understanding of deep the abyssal plain, and they encompass more than coral communities and a summary of research 30 major volcanic peaks extending from Georges priorities that could help fill these gaps are Bank southeast for about 1,100 km to the eastern highlighted as a key need to assist in identifying end of the Bermuda Rise (Figure 5.1). Several future actions to conserve and manage these recent surveys have taken place here, although vulnerable ecosystems. most of the seamount chain is located outside the 200 nautical mile U.S. Exclusive Economic Zone However, it should be noted that to our knowledge (EEZ); however, Bear, Physalia, Mytilus, and the northeast region does not have the abundance Retriever Seamounts do occur within the EEZ. of large, structure-forming deep corals and deep coral habitats that are present in other regions. In the U.S. Northeast Shelf Large Marine Thus, we will confine our discussions to the Ecosystem, deep corals have been noted since major gorgonian species such as, for example, the surveys of Verrill in the 19th century (Verrill Paragorgia arborea, Primnoa resedaeformis, 1862, 1878a, 1878b, 1879, 1884) and as fisheries Acanthogorgia armata, and Paramuricea bycatch since that period. They also occur off grandis, and others, as well as some of the Atlantic Canada on the continental slope and in more noteworthy alcyonaceans (soft corals) and submarine canyons, and are particularly abundant scleractinians (stony corals). in the Northeast Channel (Verrill 1922; Deichman 1936; Breeze et al. 1997; MacIsaac et al. 2001; II. GEOLOGICAL SETTING 1NOAA Fisheries Service, Northeast Fisheries Science Center, This brief review of the pertinent geological James J. Howard Marine Sciences Laboratory, characteristics of the regional systems of the Highlands, NJ 07732 U.S. Northeast Shelf Large Marine Ecosystem, as well as the subsequent section on the 2 New England Fishery Management Council, oceanographic characteristics, is based on 50 Water Street, Mill 2, Newburyport, MA several summary reviews. Literature citations 01950 are not included because these are generally

195 196 NORTHEAST protect thesensitive habitatstherein,including deepcoralandother structure-formingorganisms. gillnetting while onamonkfish( sidered EFH Closures andwereimplemented intheMonkfishFMP in2005.Withinthese areas,trawlingand southern NewEngland,Lydonia andOceanographerCanyonsonthecontinental shelf-breakarealsocon of MaineEssentialFishHabitat Closed Areas. In addition totheEFHClosed Areas onGeorges Bankandin exception ofthesliverthat overlapsboththeNEFMCWestern GulfofMaineGroundfish and Western Gulf NEFMC website).Nofishing restrictionsexistwithintheboundariesofStellwagen BankNMSwiththe in portionsoftheclosedareas definedintheSpecial AccessProgramduringcertain timesoftheyear(see All GearCapableofCatchingGroundfish”prohibitthe useofthegearcapablecatchinggroundfishexcept bottom-tending gearindefinitely. NortheastMultispecies Closed Areas,indicatedinthelegendby“Closuresto occurred sincethepointwassampled.EssentialFish Habitat(EFH)Closed Areas prohibittheuseofmobile Also, certaindeepcoralpointsmaynolongerbepresent duetonaturalandhumaninducedimpactsthathave tions inthisregionastheU.S.ExclusiveEconomicZone hasnotbeenextensivelysurveyedfordeepcorals. ranging from(butnotinclusiveof)1870tothe1990s. This informationdoesnotrepresentalldeepcoralloca (2003) databasesrepresentknownlocationsofstony, gorgonian,andtruesoftdeepcoralspeciesfromdata Figure 5.1 REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Coral Distribution-Deepcoraldatafromthe Theroux andWigley(1998)theWatling et al. Lophius americanus , goosefish) days-at-sea (DAS)isprohibited indefinitelyto - - STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION accepted descriptions of the regional systems. (Nantucket Shoals) to Nova Scotia (Scotian Source references include, and more information Shelf) and covers more than 40,000 km2. It is can be found in: Backus and Bourne (1987), characterized by a steep slope on its northern Schmitz et al. (1987), Tucholke (1987), Wiebe edge and a broad, flat, gently sloping southern et al. (1987), Cook (1988), Stumpf and Biggs flank. It is separated from the rest ofthe (1988), Townsend (1992), Conkling (1995), continental shelf to the west by the Great South Brooks (1996), Sherman et al. (1996), Beardsley Channel. The central region of the Bank is et al. (1997), Dorsey (1998), Packer (2003), shallow. Bottom topography on the eastern Bank Stevenson et al. (2004), Auster et al. (2005), and is characterized by linear ridges in the western Babb (2005). shoal areas; relatively smooth, gently dipping sea floor on the deeper, eastern most part, and Gulf of Maine steeper and smoother topography incised by The Gulf of Maine is a semi-enclosed continental submarine canyons on the southeastern margin. shelf sea of 90,700 sq. km bounded on the east by The sediments vary widely, but are mostly sandy

Browns Bank, on the north by the Scotian Shelf, or silty, with coarse gravel and boulders at the NORTHEAST on the west by the New England states, and on northern margins. the south by Cape Cod and Georges Bank. It is distinct from the Atlantic, an ecologically separate At least 70 large submarine canyons occur near sea within a sea. The Gulf is characterized by the shelf break along Georges Bank and the Mid- a system of 21 deep basins, moraines, rocky Atlantic down to Cape Hatteras, cutting into the ledges, and banks, with limited access to the slope and occasionally up into the shelf as well. open ocean. Sediments in the Gulf of Maine The canyons are typically “v” shaped in cross are highly variable and, when coupled with the section, and include features such as steep walls, vertical variation of water properties found in the exposed outcroppings of bedrock and clay, and Gulf, results in a great diversity of benthic habitat tributaries. Most canyons may have been formed types and benthic organisms. Sand, silt, and by mass-wasting processes on the continental clay are found throughout the Gulf, with the finer slope; some, like the Hudson Canyon (Mid- sediments generally found in the deeper basins. Atlantic), may have formed because of fluvial Rocky substrates (which include gravel, pebbles, drainage. The canyons exhibit a more diverse cobbles, and boulders) are found primarily in fauna, topography, and hydrography than the the Northeast Channel, with other smaller, more surrounding shelf and slope environments. The variable rocky areas interspersed in the Gulf. diversity in substrata types tends to make the Rocky outcrops form significant features, such canyons biologically richer than the adjacent as Cashes Ledge, and benthic fauna found on shelf and slope. these include sponges, tunicates, bryozoans, and hydroids. As mentioned above, the New England Seamount chain extends southeast of Georges Bank for Of the 21 deep basins, Jordan (190 m), Wilkinson about 1,100 km, rising as much as 4,000 m above (190 m), and Georges Basins (377 m), are the the Sohm Abyssal Plain. Of the four seamounts largest basins and deepest habitats within the within the U.S. EEZ, Bear Seamount is the closest Gulf of Maine. Their great depths resulted from and rises from a depth of 2,000-3,000 m to a glacial erosion of relatively soft rocks. The bottom summit that is 1,100 m below the sea surface. sediments of these deep basins are generally very The minimum depths of the others are: Physalia fine, featureless muds, but some gravel may also (1,848 m), Mytilus (2,269 m), and Retriever be found; little or no sediment transport occurs (1,819 m). Several other seamounts outside the here. Unique invertebrate communities are found U.S. EEZ are biologically significant because on the seafloor, including deep corals, ophiuroids they rise to relatively shallow depths. Substrate (brittle stars), tube building amphipods, burrowing types range from solid basalt to manganese anemones, and polychaete worms. crusts to rock and coral rubble to mixtures of basalt pebbles and sand to fine carbonate oozes. Georges Bank Sediments cover the summits with more exposed Georges Bank is a shallow (3-150 m depth), rock surfaces on the sides. elongate extension of the continental shelf that extends from Cape Cod, Massachusetts

197 198 NORTHEAST ae etrn truh h Nrhat Channel Northeast the through the entering water in slope important saline and warm relatively particularly Dense, spring. is coastal which from runoff rivers, freshwater and Channel, that water Northeast the through and north shelf the from enters Scotian cold fresh, by driven the Gulf along the shore. This current is primarily of margin the around flows that current surface The Gulf has a general counterclockwise nontidal Ocean. Atlantic and Maine of Gulf the between is one of the primary avenues for water exchange and Browns Bank leads into Georges Basin, and Bank Georges between Channel Northeast The Gulf ofMaine III. abundance ofinfaunalorganismsfoundthere. high the for account to helping above, shelf the off funneled are that nutrients and sediment of Cape fluxes off high exceptionally receives slope also Hatteras continental The slope. fines the 70-100% on are sediments and line,” “mud the called sometimes is which break, shelf the at rapidly increases content sediment Fine valley. Hudson the in common is but shelf, the of most over rare is Mud shelf. outer the on and valley shelf Hudson the in clay and silt, sand, fine with grains, coarse to medium mostly are sands The m. 0-10 from thickness in Sand and vary deposits gravel mud deposits. with and sand, silt and fine-grained gravel of coarse-grained of composed patches isolated primarily are and distributed, uniformly fairly are shelf Mid-Atlantic the over Sediments Hatteras. Cape off geology and oceanography unique and the and Wilmington), Washington, Baltimore, (Norfolk, Cape and Charles May Cape other between canyons several major significant include Other features slope. physiographic isobath, continental m the 40 the onto about at Shelf shelf, inner- continental the Hudson from extending the Canyon, is and Valley Mid-Atlantic the within scarps, massifs, and sand ridges. The shoal main physiographic feature channels, and shelf valleys include shelf the of features primary The canyons. submarine near except break, shelf to shore from linearly increases depth as smooth, relatively is topography shelf the Here Stream. Bank south to Cape Hatteras, and east to the Gulf Georges from waters slope and shelf continental the of region the to refers Bight Mid-Atlantic The Mid-Atlantic Bight REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF OCEANOGRAPHIC SETTING mixed vertically. This results in a tidal front that front tidal a in results This vertically. mixed well Bank very the over be waters the can keep and Bank strong, the of top shallow the over which can all occur simultaneously. Tidal currents and currents, storm-induced southeast, intermittent strong, very and northwest flow predominantly tidal around semidiurnal strong gyre a clockwise Bank, Georges persistent a is There Georges Bank salinity bottomwater. high- cold, cool and water; water; intermediate nutrient-rich surface nutrient-poor warm, into Wilkinson, and Georges Basins becomes Jordan, layered in water the summer, the In variations. moving currents, and annual and seasonal inflow Stream rings, strong winds which can create fast Gulf by water shelf of entrainment the as such interactions shelf-slope to due year to year from significantly vary can circulation Gulf formation. gyre influences also slope continental the from many physicalfactors. the by influenced be can and of variable, highly is front position The offshore. further km 25-55 surface. It reaches surface waters approximately the toward east the to up slopes water,then and of depth m 75-100 about at bottom touches and shelf the of edge the at located usually is front This front. shelf-slope the called is meet masses water. The abrupt gradient shelf where these than two saline water more and water warmer be Slope to tends Stream. Gulf the from meanders or rings core warm is by that interrupted flow occasionally southwestward slow a have Mid-Atlantic Bight the of waters slope and shelf The Mid-Atlantic Bight Limanda ferruginea cod), and, productivity includes such species as primary which production, of fish of levels high historically, levels high by characterized is and conditions environmental many community, by influenced is that community biological planktonic the including larval fish. Georges Bank has of a diverse distribution in instrumental is gyre clockwise The Bank. the of shelf waters well-mixed on the seaward and shoreward stratified sides the seasonally warmer, of the from shallows waters cool the separates Melanogrammus aeglefinus (yellowtailflounder). Gadus morhua (haddock), and (Atlantic STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

IV. STRUCTURE AND HABITAT-FORMING a complete list of the deep coral species in this DEEP CORALS region, see Appendix 5.1.

As stated in the introduction, the northeast region a. Stony Corals (Class Anthozoa, Order does not appear to have an abundance of large, Scleractinia) structure-forming deep corals and deep coral Cairns and Chapman (2001) list 16 species habitats. Thus, we will confine our discussions of stony corals from the Gulf of Maine and primarily to the major deep gorgonian corals as well Georges Bank to Cape Hatteras (Table 5.1) as to some of the more noteworthy occurrences (See also Cairns 1981). Most of the stony corals of alcyonaceans (soft corals) and scleractinians in this region are solitary organisms and one (stony corals), including solitary species or species, Astrangia poculata, can occur in very those found in shallower habitats. Records of shallow water, at depths of only a few meters. gorgonians and soft corals are often combined Moore et al. (2003, 2004) reported several into one database, so for convenience we will be species of solitary and colonial stony corals on discussing those two orders simultaneously. For Bear Seamount; one notable solitary species, NORTHEAST

Table 5.1. Stony corals from the Gulf of Maine and Georges Bank to Cape Hatteras, based on Cairns and Chapman (2001). Vaughanella margaritata has been reported from Bear Seamount of the New England Seamount chain (Moore et al. 2003). Depth ranges are for the western Atlantic only. * = potentially structure-forming colonial species. Taxon Distribution Coloniality Attachment Depth (m) Astrangia poculata (Ellis & Endemic to western Atlantic colonial attached 0-263 Solander, 1786) Caryophyllia ambrosia Widespread (cosmopolitan) solitary free-living 1487-2286 ambrosia Alcock, 1898 distribution Caryophyllia ambrosia Endemic to western Atlantic solitary free-living 183-1646 caribbeana Cairns, 1979 Dasmosmilia lymani Widespread (cosmopolitan) solitary free-living 37-366 (Pourtales, 1871) distribution Deltocyathus italicus Amphi-Atlantic with a solitary free-living 403-2634 (Michelotti, 1838) disjunct distribution Desmophyllum dianthus Widespread (cosmopolitan) solitary attached 183-2250 (Esper, 1794) distribution Enallopsammia profunda Endemic to western Atlantic colonial attached 403-1748 (Pourtales, 1867)* Enallopsammia rostrata Widespread (cosmopolitan) colonial attached 300-1646 (Pourtales, 1878)* distribution Flabellum alabastrum Amphi-Atlantic with solitary free-living 357-1977 Moseley, 1873 contiguous distribution Flabellum angulare Moseley, Amphi-Atlantic with solitary free-living 2266-3186 1876 contiguous distribution Flabellum macandrewi Gray, Amphi-Atlantic with solitary free-living 180-667 1849 contiguous distribution Fungiacyathus fragilis Sars, Widespread (cosmopolitan) solitary free-living 412-460 1872 distribution Javania cailleti (Duch. & Widespread (cosmopolitan) solitary attached 30-1809 Mich., 1864) distribution Widespread (cosmopolitan) Lophelia pertusa (L, 1758)* colonial attached 146-1200 distribution Solenosmilia variabilis Widespread (cosmopolitan) colonial attached 220-1383 Duncan, 1873* distribution Vaughanella margaritata Endemic to northwestern solitary attached 1267 (Jourdan, 1895) Atlantic

199 200 NORTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF l (90 18) se ecitos eo] Yale et below], descriptions see Hecker 1983); (1980, al. (1980), Blechschmidt and 1990), Hecker (1980, Hecker Hecker [e.g.; soft (1936), collaborators and true Deichmann sources, Verrill, of and variety including a gorgonian from occurrences both coral of obtained database (2003) records al. et Watling The OrderGorgonacea) d. Gorgonians( have beenfoundinthisregion. order this from species of records No c. observe ontheseamounts. previously not did they that species 7 including Seamounts, Rise England Corner and New the to expedition 2005 their during coral black of species 15 and Brugler collected and observed (2006) France expedition; 2004 their of black coral from the seamounts during et al. (2005) collected at least 8 species Albatross R/V the by 1883 in collected also was and collections, Institution Smithsonian Cirrhipathes 2005). (Brugler sp. coral, black Another species on living found associated been have few Seamount very England chain, New the Within Bank. Georges off area same the near in collected 1883 by the R/V been having Smithsonian Institution, the of collections the in Bear Seamount (Brugler on 2005); m it is 1643 also from found Atlantic. collected been northwest the ( coral black Bushy in occur predominantly to known are species some but are tropical, Antipatharians b. Watling etal.2005;Shank2006). 2006; al. et (Adkins corals stony found also have Seamounts Rise Corner and Other England New the to 2001). expeditions recent Chapman and (Cairns and is endemic to the northwest Atlantic original description over 100 years ago, its since species this of record first the margaritata Vaughanella Gold Corals Black Corals Order Zoanthidea) Order Antipatharia) at 262 m off Virginia. WatlingVirginia.off m 262 at p, s lo on i the in found also is sp., (Class Anthozoa, (Class Anthozoa, Albatross Class Anthozoa, Leiopathes represents , from 1754 m Leiopathes sp.) has sp.)

tion, Washington, D.C.,pers.comm.). (ITIS 2006)database/websiteandS.Cairns(SmithsonianInstitu changes basedonIntegrated Taxonomic InformationSystem Cape Hatteras,fromWatling and Auster (2005),withtaxonomic on thenortheasternU.S.continentalshelfandslopenorthof Table 5.2. Primnoidae Chrysogorgiidae Paragorgiidae Anthothelidae Paramuriceidae Acanthogorgiidae Order Gorgonacea Nephtheidae Alcyoniidae Clavulariidae Order Alcyonacea Isididae Taxa List ofgorgoniansandsoftcoralsknowntooccur aauiede Atohlde Paragorgiidae, Anthothelidae, Paramuriceidae, (Acanthogorgiidae, families gorgonian seven the in species 17 These (Table5.2). Hatteras Cape for recorded the northeastern U.S. shelf and slope north of were families 7 in species 17 of total A (2005)]. Auster and Watling see information, Undersea National Research Center (NURC) field studies recent [for further from NEFSC and taxa, observations coral the identified of database collections, benthic museum Peabody Narella Radicipes gracilis Iridogorgia pourtalesii Chrysogorgia agassizii Paragorgia arborea Anthothela grandiflora Swiftia Paramuricea Paramuricea Paramuricea Acanthogorgia armata Capnella Capnella Gersemia Gersemia rubriformis Anthomastus agassizii Anthomastus grandiflorus Alcyonium digitatum Clavularia rudis Clavularia modesta Lepidisis Keratoisis Keratoisis Acanella Thouarella Primnoa

casta laxa

resedaeformis arbuscula caryophyllia glomerata florida

fructicosa grayi ornata grasshoffiCairns,2006 Deichmann,1936 (Verrill, 1883) n.sp. placomus grandis Wright, 1869 (Verrill, 1922) (Rathke,1806) Species Verrill, 1878 (Verrill, 1884) (Johnson,1862) (Verrill, 1874) (Linné,1758) (Verrill, 1869 (Sars,1860) Linné,1758 Verrill, 1883 (Ehrenberg,1934) Verrill, 1883 Verrill, 1883 (Sars,1856) Verrill, 1878 Verrill, 1922 (Verrill, 1883) (Linné,1758) (Gunnerus,1763) Verrill, 1878 - STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION Reference U.S. NMNH collection, OBIS U.S. NMNH collection, OBIS U.S. NMNH collection, OBIS U.S. NMNH collection U.S. NMNH collection U.S. NMNH collection, OBIS U.S. NMNH collection, OBIS U.S. NMNH collection, OBIS U.S. NMNH collection, OBIS U.S. NMNH collection U.S. NMNH collection, OBIS U.S. NMNH collection U.S. NMNH collection, OBIS U.S. NMNH collection U.S. NMNH collection, OBIS U.S. NMNH collection U.S. NMNH collection, OBIS NORTHEAST Depth Range (m) 274-3651 430-2491 (1538 m min in NE U.S.) 393-2199 (1330 m min in NE U.S.) 1977-2249 2491-4332 119-3316 1850-2140 219-2295 (doubtful report at 59 m) 1211-2844 1483-2359 1334-2194 2513-4332 1502-2505 2683-3740 (3166 m min in NE U.S.) 549-3338 (1538 m min in NE U.S.) 37-2249 (229 m min in NE U.S.) 20-812 (51 m min in NE U.S.) Distribution Newfoundland to Antarctica, N. Europe Chile, Hawaii, Bahamas, Lousiana, Lydonia Canyon to Puerto Rico, Africa, N. Europe Aleutians, Japan, W. Hawaii, Newfoundland to South Carolina, Japan, Africa, N. Europe W. Nova Scotia to Virginia Massachusetts to Virginia Iberia, California, Virginia, to Newfoundland Africa N. Bay of Biscay New Jersey, Newfoundland to North Carolina, California N. Africa, W. Carolina, North to Scotia Nova Europe Nova Scotia to Virginia Africa, W. Carolina, North to Massachusetts N. Europe Massachusetts to Virginia Panama, Carolina, North to Massachusetts Africa W. Louisiana Massachusetts to Virginia, Massachusetts to Louisiana, Suriname, N. Europe, Indian O. the Virgin Newfoundland Islands, to Alaska Islands, (doubtful), the Virgin Massachusetts, NC Iberia York-Florida, New Species Anthoptilum grandiflorum Anthoptilum murrayi Kophobelemnon stelliferum Kophobelemnon scabrum Kophobelemnon tenue Pennatula aculeata Pennatula grandis Pennatula borealis Distichoptilum gracile Protoptilum abberans Protoptilum carpenteri Scleroptilum gracile Scleroptilum grandiflorum Umbellula guntheri Umbellula lindahlii Balticina finmarchica Stylatula elegans Identity and distribution of Pennatulacea on the northeastern U.S. continental shelf and slope. NMNH = Smithsonian Natural Museum of Natural Higher Taxon Anthoptilidae Kophobelemnidae Pennatulidae Protoptilidae Scleroptilidae Umbellulidae Virgulariidae Table Table 5.3. History; OBIS = Ocean Biogeographic Information System.

201 202 NORTHEAST Moe t l 20, 04 Wtig t al. et 2005, 2006). Watling 2004; 2003, al. et (Moore New England and Corner Rise Seamounts the to expeditions recent during collected been also have interpreted Gorgonians caution. be with should surveys these so various surveys (Watling and Auster 2005), these from identifications the of accuracy the about uncertainty is there reasons, of Auster and the 2005). It should be noted that, for a variety (Watling of slope waters deeper continental the in of most abundant being as because well as sizes, documented larger their best the are Chrysogorgiidae, Primnoidae, and Isiddae) STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Wigley (1998)database. the Mid-Atlantic.Sourcecredit: Theroux and ( Bank. B)Distributionofstonyorhardcorals bellum corals ( Figure 5.2 Astrangia ) intheGulfofMaineandGeorges Astrangia, Dasmosmilia , A Dasmosmilia . Distributionofstonyorhard , and A Flabellum , and Fla - ) in B common in the database, but apparently but database, the in not common were that species Two 5.2). (Table Hatteras Cape of north slope and shelf U.S. northeastern the for recorded were Auster and families 3 in species 9 of total WatlingA (2005)]. see information, further [for studies field (NURC) Center museum Research Undersea National recent from Peabody observations and taxa, coral identified of Yale database benthic NEFSC the collections, descriptions see 1983); below], (1980, al. et Hecker andBlechschmidt(1980), including 1990), and (1980, Hecker [e.g.; collaborators sources, (1936),Hecker of Deichmann Verrill, variety both a from of records gorgonian and true soft coral occurrences obtained database As stated above, the Watling et al. (2003) Order Alcyonacea) e. True SoftCorals t hud e oe ta, o a variety a for that, noted be should gorgonians, the it with As 2005). Auster and corals soft the were records, nearshore in numerous very are Alcyonium pce (aln and (Watling species Gersemia rubiformis Gersemia (Class Anthozoa, STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION of reasons, there is uncertainty about the accuracy of the identifications from these various surveys (Watling and Auster 2005), so these surveys should A be interpreted with caution. Soft corals have also been collected during recent expeditions to the New England and Corner Rise Seamounts (Moore et al. 2003; Watling et al. 2005, 2006). f. Pennatulaceans (Class Anthozoa, Order Pennatulacea) Records of sea pens were drawn from Smithsonian Institution collections and the Theroux and Wigley benthic database. NORTHEAST Nearly all materials from the former source were collected either by the U.S. Fish Commission (1881-1887) or for the Bureau of Land Management (BLM) by the Virginia Institute of Marine Sciences (1975-1977) and Battelle (1983-1986). These latter collections heavily favor the continental slope fauna. The Theroux and Wigley collections (1955-1974) were made as part of a regional survey of all benthic species (Theroux and Wigley

Figure 5.3 A) Distribution of gorgonians and soft corals in the Gulf of Maine and Georges B Bank. B) Distribution of gorgonians and soft corals in the Mid-Atlantic. Source credits: Ther- oux and Wigley (1998) database, Watling et al. (2003) database.

1998), heavily favoring the continental shelf fauna. A list of 17 sea pen species representing five families was compiled from these sources for the northeastern U.S. (Table 5.3). None commonly occur in shallow water, and only two species are known from the lower continental shelf depths (80-200 m) in this region: Pennatula aculeata (common sea pen) and Stylatula elegans (white sea pen). P. aculeata is common in the Gulf of Maine (Langton et al. 1990), and there are numerous records of Pennatula sp. on the outer continental shelf as far south as the Carolinas in the Theroux and Wigley database. S. elegans is abundant on the Mid-Atlantic coast outer shelf (Theroux and Wigley 1998). The other 15 sea pen species have been reported exclusively from the continental slope

203 204 NORTHEAST been foundinthisregion (Cairns1992). have family this from species of records No g. sediment whendisturbed. the into of colony entire the or capable part retracting buried are species swollen, Some a peduncle. by place in anchored sediments, muddy in other live pens sea most corals, Unlike m). (200-4300 depths Source credit:Watlingetal.(2003)database and Acanthogorgia armata,Paragorgiaarborea, Georges Bank.B)Distributionofthegorgonians Primnoa resedaeformis Acanthogorgia armata,Paragorgiaarborea Figure 5.4 STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Primnoaresedaeformis Stylasterids Family Stylasteridae) Anthothecatae, SuborderFilifera, A A) Distributionofthegorgonians (Class Hydrozoa,Order in theGulfofMaineand intheMid-Atlantic. , and . B and B, respectively. There appears to be to appears There respectively. B, and and 5.2A Figures Bank, in depicted are Maine/Georges Mid-Atlantic of Gulf the in database (1998) Wigley and Theroux only Dasmosmilia specifically subclass of corals, stony the distributions the in The anemones Zoantharia. of types various the with together lumped are sp. stony corals e.g., such as groups; taxonomic between distinguish from not do often taken They 1956-1965. samples on based Atlantic, distribution of deep corals in the northwest the described (1998) Wigley and Theroux Georges Bank,andMid-AtlanticBight General DistributionintheGulfofMaine, V. and B). 5.2A (Figure margin continental the along Bank (Figure 5.2A), but note their presence Georges on corals stony of lack general a HABITATS OF CORAL SPECIES AND SPATIAL DISTRIBUTION and , Astrangia Flabellum sp. and fo the from , Astrangia Flabellum ,

STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

A B NORTHEAST

Figure 5.5 Distribution of the gorgonians Acan- thogorgia armata, Paragorgia arborea, and Prim- noa resedaeformis in A) Lydonia and B) Oceanog- rapher Canyons (Gulf of Maine) and C) Baltimore Canyon (Mid-Atlantic). Source credit: Watling et al. C (2003) database.

Theroux and Wigley (1998) also discussed the soft corals, gorgonians, as well as the sea pens. They were present along the outer margin of the continental shelf and on the slope and rise, and were sparse and patchy in all areas, particularly in the northern section. Theroux and Wigley (1998) found that they were not collected in samples taken at <50 m in depth, and were most abundant between 200-500 m. Figures 5.3A and B depict the distribution of gorgonians and soft corals in the Gulf of Maine, Georges Bank, and the Mid- Atlantic, based on the Theroux and Wigley (1998) database and the Watling et al. (2003) database. Identified species include gorgonians such as Acanella sp., Paragorgia arborea, Primnoa reseda [now resedaeformis, see Cairns and Bayer (2005)] and the soft coral Alcyonium sp. Gorgonians and soft corals were collected from gravel and rocky outcrops (Theroux and Wigley 1998).

Watling and Auster (2005) noted two distinct distributional patterns for the gorgonians and 205 206 NORTHEAST erfrne dt o te rsne f deep of corals inthis region. presence the available on best data georeferenced the represents databases these of two combination the However, remain species unknown. these of many true of the distributions identified, since not and were surveyed specimens been some not have areas all Since orcollected. observed were identified be i.e., only; could that corals deep where describe presence only they show databases, al. et (2003) Watling and the (1998) both Wigley and on Theroux based maps chapter, distribution this in the presented that noted be should It observations. (Figure (1987) Grosslein’s and Bank Theroux affirming 5.4A), Georges of Peak Northeast the on and Maine of Gulf the throughout found was addition, and In B, respectively). 5.5A, C (Figure canyons Oceanographer, Baltimore Lydonia,and from come three these species for records of majority The coast. U.S. England/Mid-Atlantic New the off canyons submarine major the of locations the shows 5.1 Canyons in the Mid-Atlantic (Figure 5.4B). Figure Norfolk and Baltimore and 5.4A), (Figure Georges Bank off located Canyons Oceanographer database. Lydoniain occur species and three All (2003) al. et Mid-Atlantic, Watling the on the based respectively, and Bank Maine/Georges of arborea P. of distribution the gorgonians: major three depict B and 5.4A Figures et al.2002). to off Virginia Beach, Virginia (37°03’N) (Heikoop (Tendal Atlantic North 1992); the in widespread are species Both Bank. Georges of Peak Northeast as well as reported (1987) Grosslein and Theroux while Maine, of Gulf the of fauna gravel by Wigley (1968) as a common component of the genus upper arborea the to gorgonians the include and waters slope continental shelf throughout occur Anthomastus and genera Acanella the in gorgoninans of species include that these m; species >500 depths at deepwater occur are Most corals. soft REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Swiftia P. resedaeformis Paramuricea , , . resedaeformis P. and , Anthothela ad ot oas n h genera the in corals soft and , . arborea P. and . resedaeformis P. . Clavularia . arborea P. , t b cmo o the on common be to , has been reported south cnhgri armata Acanthogorgia Lepidisis ad pce i the in species and , Ohr species Other . P. P.resedaeformis a described was

Acanthogorgia resedaeformis , n h Gulf the in Radicipes P. , , , ,

lca ertc ad oa rbl poie hard provide rubble coral and erratics Canyon, glacial Oceanographer of flank west the On Oceanographer Canyon et al.1983). (Hecker slope the on boulders and outcrops on found also were axis, canyon the along outcrops grandis P. in Lydonia Canyon; for example, important most the factor in determining the distribution of the corals be to appeared substrate suitable of availability The 1983). al. et (Hecker sp. include canyon the in found corals stony Other wall. areas west the some of in and axis the of part deeper the cristagalli coral stony solitary The canyon. the of walls west and east the both on occurred species, related closely a east of individuals the Several m. 500-700 up between axis farther common most was which the soft coral was found coral 1980). abundant al. most et The a major component of the fauna, and most were most and fauna, the of component major a were corals deep LydoniaCanyon, of axis the In Lydonia Canyon also included. slope areas; the surveys of continental other researchers are and canyons submarine prominent the of several for findings their of summary a is following slope The 1980). continental Blechschmidt and the (Hecker on and canyons in both canyons while the soft substrate types were found in only found were substrates, hard to restricted those as such species, some and canyons, the 1980, in al. diverse more and et denser were Corals (Hecker 1983). sled camera towed and submersible via 1980s the in U.S. northeastern the the of off canyons fauna several and margin epibenthic continental and corals deep the Dr. Barbara Hecker and her colleagues surveyed Canyons Continental Margin/SlopeandSubmarine coral and included canyon on hard substrates occurring along the axis of the gorgonians Other m. >800 depths at axis the of P. grandis armata gorgonians The restricted to hard substrates (Hecker et al. 1980). Jvna cailleti Javania , nhtea grandiflora Anthothela Trachythela were found in the axis and on the walls, with (= being more common in the deeper part wih ee omn naiat of inhabitants common were which , dianthus Eunephthya amsii lymani Dasmosmilia aauie grandis Paramuricea (now . roe, . resedaeformis P. arborea, P. and ) was found on outcrops in outcrops on found was ) Clavularia oeomla variabilis Solenosmilia (now a wl as well as , anla glomerata Capnella Capnella D. dianthus ) Desmophyllum rudis , Flabellum

and (Hecker ) h soft the florida, and A. , ,

STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION substrate for the attachment of several species of constricts and bends, outcrops and talus blocks coral, the most common being the gorgonian A. are exposed. Several corals restricted to hard armata from 650-950 m (Hecker and Blechschmidt substrates were found in this area by Hecker et 1980; Hecker et al. 1980). A. armata and P. al. (1980). Massive colonies of the gorgonian P. grandis were also found on the east flank, and on arborea were found on the large rock outcrops. both the east and west walls along the axis. The Other corals found included the gorgonians soft coral C. florida was also abundant on the east A. armata, P. resedaeformis, A. grandiflora, A. flank. The most common species in the deeper arbuscula, and the soft corals C. florida and A. zone was the gorgonian P. grandis (Hecker et agassizii (Hecker et al. 1980, 1983). The solitary al. 1980). Hecker and Blechschmidt (1980) and stony coral D. lymani occurred in dense localized Valentine et al. (1980) also noted the presence patches near the head of Baltimore Canyon, but of the Paramuricea borealis (now P. grandis), in was absent from many other areas in the Canyon Oceanographer Canyon; Valentine et al. (1980) (Hecker et al. 1983). Other stony corals found observed their greatest abundance to be from included Flabellum sp. and D. dianthus (Hecker

1100-1860 m, while Hecker and Blechschmidt et al. 1983). NORTHEAST (1980) observed that they were dominant from 950-1350 m. A. armata was most common on Other Canyons smaller cobbles, boulders, and coral rubble while Hecker and Blechschmidt (1980) surveyed the P. grandis was usually found on large boulders deep corals and epibenthic fauna of the several or outcrops (Hecker and Blechschmidt 1980; other canyons off the northeastern U.S. Discrete Hecker et al. 1980). Other deep corals on hard assemblages of corals were not identified. For a substrates in Oceanographer Canyon included complete list of species found in the historical the soft coral Anthomastus agassizii in the axis survey of Hecker and Blechschmidt (1980) and on both walls, the soft coral Clavularia and the Hecker et al. (1980) field study, see rudis on the west wall, large colonies of the Appendix 5.2, which includes Opresko’s (1980) gorgonian P. arborea in the axis above 1000 m, list of octocorals and Hecker’s (1980) list of and numerous individuals of the small encrusting scleractinians from those two surveys. gorgonian, A. grandiflora, along the axis. Valentine et al. (1980) also found A. agassizii in In Heezen Canyon, the gorgonian A. arbuscula a zone of greatest abundance from 1100-1860 and the soft coral A. grandiflorus, both found m, while Hecker and Blechschmidt (1980) found on soft substrates, occurred at 850-1050 m; the this species mostly from 950-1350 m on glacial gorgonian P. grandis was common from 1450- erratics, outcrops, and coral rubble. The solitary 1500 m; the soft coral A. agassizii and the stony stony coral, D. dianthus, was found throughout coral D. dianthus were found from 1150-1500; D. the axis between 1500-1600 m (Hecker and dianthus was also found from 1500-1550 m. The Blechschmidt 1980) and on the west flank. Deep walls of Corsair Canyon were heavily dominated corals restricted to soft substrates included the by corals, all of which were restricted to soft soft coral Anthomastus grandiflorus on the east substrates. The gorgonian A. arbuscula was flank (Hecker and Blechschmidt 1980; Hecker et prominent from 600-800 m, and the soft coral al. 1980) and the gorgonian Acanella arbuscula A. grandiflorus dominated from 800-1000 m. In on both walls (Hecker and Blechschmidt 1980; Norfolk Canyon in the Mid-Atlantic, the stony coral Hecker et al. 1980); A. arbuscula was found by D. dianthus and the gorgonian A. armata were Hecker and Blechschmidt (1980) mostly from found on hard substrate at 1050-1250 m; both 950-1350 m. The gorgonian P. resedaeformis is were also observed in this canyon by Malahoff et also found in the canyon, with a zone of greatest al. (1982). Hecker and Blechschmidt (1980) also abundance from 300-1099 m (Valentine et al. noted that the solitary stony coral Flabellum sp. 1980). was seen in high concentrations at 1300-1350 m depth in Norfolk Canyon, and the soft coral A. Baltimore Canyon grandiflorus was found between 2150-2350 m. Compared to Lydonia and Oceanographer Canyons, Baltimore Canyon in the Mid- Deep corals have been seen on the shelf around Atlantic had the fewest corals, perhaps due to Hudson Canyon and in the head of the Canyon the scarcity of exposed outcrops (Hecker et al. (see Appendix 5.2). For example, most recently 1980). At depths >400 m, where the canyon axis a survey by Guida (NOAA Fisheries Service,

207 208 NORTHEAST ekr t l (93 sree a ae called area an Slope III, a 25 surveyed mile wide section of the (1983) continental al. et Hecker Other Areas of theContinentalSlope of thecanyon. of Gersemia populations coral soft abundant the Blechschmidt found and who Hecker (1980), from comes canyon itself the within deep occurring corals deep lower.much were However,of evidence only the exceeded 200 polyps m patches those within densities local m; 105-120 along the canyon’s rim near its head at depths of particularly abundant in patches in a narrow band the at depths of 100 to found 200 m (Figure 5.6). They were 2004 and solitary stony coral 2002, 2001, in Canyon Hudson around shelf the on habitats data) benthic of unpublished Sciences NJ, Marine Highlands, Howard Laboratory, J. James NEFSC, STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF August 2004,andMarch2007. Data obtainedfromstillphotosandtrawlsamplestakenduringOctoberNovember2001,2002,2005, Dasmosmilia lymani Figure 5.6.

fructicosa Distribution andapproximatedensities(polypspersquaremeter)ofthesolitarystonycoral uehha fruticosa Eunephthya D. lymani in samplesfromtheMid-AtlanticshelfaroundHudsonCanyon(Guida,unpublisheddata). ?)

only in the deeper portion deeper the in only -2 , but densities elsewhere at a number of sites sm as (same both slopes. gorgonians The as well Flabellum included I Slope on found corals stony Other m. <500 depths at areas slope both in abundances solitary stony coral gorgonian the corals and stony solitary the included III Slope on found corals Deep north. the to Canyon by bounded TomsMeys and south the to Canyon was and I, Slope of north miles 70 about was II, Carteret Slope other,called the and and north, the on Canyon south the on slope Canyon Lindenkohl two by flanked was surveyed I, Area Slope called they one, areas; Mid-Atlantic Canyons. the Hydrographer In and Veatch between slope on the southwestern edge of Georges Bank, h sf coral soft the D. dianthus D. S. variabilis S. sp.

. agassizii A. and P.grandis , the soft coral soft the , P.grandis D. lymani , were also found on Slope II. Slope on found also were , . dianthus D. . In the Mid-Atlantic, the Mid-Atlantic, the In . ee lo rsn on present also were and occurred in very high toe oas as corals, those ; A. agassizii A. A. arbuscula A. . lymani D. , and , and

STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

Hecker (1990), in a later survey of the megafaunal and the colonial species Lophelia pertusa. assemblages at four locations on the continental Desmophyllum cristagalli (= dianthus) had been slope south of New England [including two that found on the Seamount during previous surveys. were surveyed as part of the Hecker et al. (1983) Gorgonians found by Moore et al. (2003) included study] found that the solitary stony corals D. lymani Paragorgia sp., Lepidisis sp. Swiftia (?) sp., and and Flabellum sp. dominated the fauna on the Acanthogorgia angustiflora, although the latter upper slope, although D. lymani was absent from species does not appear anywhere else in the their transect off Georges Bank. The gorgonian deep coral records from this region. Although A. arbuscula and the soft coral C. florida, which Moore et al. (2003) mentions that a species of dominated the fauna on the shallower section of Primnoa had been found on the Seamount during the middle slope, were found only at their transect previous surveys, Cairns (Smithsonian Institution, off Georges Bank; the soft coral A. agassizii was Washington, D.C., pers. comm.) states that also found in dense populations there. Primnoa does not occur on Bear Seamount; this record had originally been taken from Houghton The New England Seamounts et al. (1977), which is in error. Moore et al. (2003) NORTHEAST Deep corals are one of the dominant members also collected the soft coral A. agassizii. of the epifaunal communities on the New England Seamount chain (Auster et al. 2005). Deep coral collected on Bear Seamount during A 2004 exploratory cruise to the New England a follow-up cruise in 2002 by Moore et al. (2004) Seamount chain revealed significant deep coral included the solitary stony corals Caryophyllia assemblages, with 27 octocoral species, 8 black cornuformis (this species is not mentioned coral species, and 2 stony coral species collected elsewhere) and F. alabastrum and the colonial (Watling et al. 2005), including possible new stony corals S. variabilis and Enallopsammia species of gorgonians from the Chrysogorgiidae rostrata. Gorgonians collected included and Paragorgiidae families (Eckelbarger and Paragorgia sp., Paramuricea sp., Keratoisis sp., Simpson 2005). A 2005 cruise to the New Swiftia pallida (this species is not mentioned England and Corner Rise Seamounts sampled elsewhere), Lepidisis sp., and Radicipes gracilis. 39 species of octocorals, including 7 that may be new to science, and observed and collected 15 species of black coral, including 7 species that VI. SPECIES ASSOCIATIONS WITH DEEP the researchers have not previously observed CORAL COMMUNITIES on the seamounts (Brugler and France 2006; Watling et al. 2006). Distributions of several Commercial Fisheries Species species (e.g., Paragorgia sp., Lepidisis spp., The role of deep corals as possible habitats for Paramuricea spp., as well as stony corals and fishes has only recently been addressed in the black corals) are currently being quantified using literature. The corals Primnoa, Lophelia, and videotapes and digital still images (Figure 5.7). Oculina from other regions have been the most Preliminary quantitative analyses of coral species studied. Several studies have documented that distributions indicate that community composition certain fish commonly occur in the vicinity of differs considerably between seamounts, corals more often than in areas without corals. even at comparable depths. These differences In the northwest Atlantic, this has been noted correspond to biogeographical boundaries, or for Sebastes sp. (redfish) in the Northeast they may be due to species’ responses to local Channel (Mortensen et al. 2005). Redfish may habitat conditions, such as substratum type or take advantage of structure on the bottom as a flow. Substantial variation in faunal composition refuge from predation, as a focal point for prey, occurs between sites on a single seamount (P. and for other uses. However, in a survey of Auster, pers. comm.). habitats in the Jordan Basin in the Gulf of Maine containing coral assemblages (primarily from the During surveys of Bear Seamount within the genera Paragorgia, Paramuricea, and Primnoa), EEZ during 2000, Moore et al. (2003) noted Auster (2005) found that densities of redfish the presence of several species of stony were not significantly different between dense corals, including the solitary corals Caryophyllia coral habitats and dense epifauna habitats, ambrosia, Flabellum alabastrum, and, as although the density of redfish in these two discussed previously, Vaughanella margaritata habitats was higher than in the outcrop-boulder

209 210 NORTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF tutrl trbts f aia, u my o be not may but habitat, of attributes important structural provide do corals The communities. support high density, and unique or high diversity distribution the on of abundance effect some corals have deep do that suggests (2005) Auster not. are epifauna dense containing habitats cobble boulder- but 2003), al. Watling5.2Aet 5.3A; and (Figs. Maine of Gulf the in rare fairly are habitats coral Deep communities. and species particular of demography the in play habitats such role the that type throughout a region that will ultimately determine habitat each of states distribution actual the (2005) is it Auster corals, containing habitat a to fish of densities similar support can corals deep without habitat a that this shows study While epifauna. sparse containing habitat can affect how they will be used by fishes. For fishes. by used be will they how affect can of the deep corals themselves within a landscape forms morphological the in variations addition, In other denseepifaunalassemblages. by provided structures than different functionally A C shes, but by themselves do not do themselves by but fishes, fish example, on the New England Seamounts, Auster coral grandis as such gorgonians largest the on shrimp of occurrence al. et Hecker U.S., northeastern the Off 2000)]. (Mortensen Norway 2002); Wing and (Krieger [e.g., regions other invertebrates from and in corals deep species corals the of about associations deep known is with more region; this associations species invertebrate about available data few are There Invertebrates polyps themselves. coral, or perhaps they were foraging on the coral the with associated prey for spot foraging a as and flow from refuge a as them utilizing perhaps with associated be to appeared observe one fish species, did (2005) al. et 2005). Auster al. et (Auster flow from refuge or shelter different of terms a in value have habitat may latter the of density and form The landscape. the across spread are that branching” density high with fans robust wide) (e.g., gorgonians the summits, seamount near the on while and stands, dense moderately species forming taller the with branching,” density with fans low short and species of whip-like amixture and “tall as flanks seamount the on deep et al. (2005) describes the gorgonian assemblage Stones Science Team, IFE,URI,andNOAA-OE Lepidisis leafy sponge(centerright).B) tom center),brisingidseastar(centertop)and ( Seamount, nearthesummit. A) Bamboocoral in coralcommunitiesfromtheupperslopeofBear Figure 5.7. Keratoisis Paragorgia anla florida Capnella , sp.Photocredit:Deep Atlantic Stepping rmo resedaeformis Primnoa sp.)atleftwithglobularsponge(bot Photographicexamplesofvariations aaoga arborea Paragorgia p) r otn lre ~ m (~1 “large often are sp.)

18) oe te frequent the noted (1980) A ohuod n an and ophiuroid An . Neocyttus helgae Lophelia Primnoa Keratoisis , Paragorgia ad h soft the and , Paramuricea

f Alaska off sp.C) pertusa - B , that sp., off

STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION anemone appeared to have an association, al. 2002), and recruitment rates may also be low possibly obligatory, as suggested by Hecker et (Krieger 2001), which makes their recovery from al. (1980), with Paramuricea grandis. Hecker disturbances difficult over short time periods. Of and Blechschmidt (1980) also noted several the various fishing methods, bottom trawling has faunal associations in their study. The ophiuroids been found to be particularly destructive (Rogers Ophiomusium lymani and Asteronyx loveni were 1999; Hall-Spencer et al. 2001; Koslow et al. often associated with corals; e.g., the latter with 2001; Krieger 2001; Fosså et al. 2002; Freiwald the gorgonian A. arbuscula. Pycnogonids were 2002). seen on the gorgonian Paramuricea grandis. The effects of current and historic fishing Current studies are looking at the species efforts on deep coral and coral habitats in the associations of the octocorals of the New northeastern U.S. have not been quantified. The England and Corner Rise Seamounts. Ophiuroids types of fishing gear used here include fixed gear and marine scale worms were found to live such as longlines, gillnets, and pots and traps, commensally in and on the octocorals found on as well as trawls and dredges. Fixed gear can be NORTHEAST the seamounts (Watling 2005; Watling and Mosher lost at sea, where they can continue to damage 2006); specimens of octocorals, stony corals, corals. In Canada, longlines have been observed and their associated species from the seamounts entangled in deep corals such as Paragorgia and are also being used in population genetic studies Primnoa and may cause breakage (Breeze et that will assess, for example, the taxonomic al. 1997; Mortensen et al. 2005). Further, “ghost relationships of the corals and associated species fishing” of nets entangled in coral reefs has been (Shank et al. 2006). Within the EEZ, Moore et al. reported in the North Sea (ICES 2003). Bottom (2003) noted the following invertebrates found in trawling was found to have a larger impact on direct association with Lophelia petusa trawled deep corals per fishing unit of effort compared from Bear Seamount: a large polychaete worm to longlining; e.g., damage to Primnoa off Alaska living in tubular spaces within the coral colony, (Krieger 2001). The northeastern U.S. fisheries the attached solitary stony coral, Caryophyllia that have the highest likelihood of occurring near ambrosia, small serpulid worm tubes, a thecate concentrations of known deep coral habitats hydroid, and the gorgonian Swiftia. (e.g., in canyon and slope areas) are the Lophius americanus (monkfish or goosefish) and Lopholatilus chamaeleonticeps (tilefish) fisheries, VII. STRESSORS ON DEEP CORAL and the Chaceon (Geryon) quinquedens (red COMMUNITIES crab) and offshore Homarus americanus (lobster) pot fisheries. Fishing Effects Deep corals provide habitat for other marine Effects of other human activities life, increase habitat complexity, and contribute to marine biodiversity, and their destruction Invasive Species could have a significant impact on other marine Little is known about any existing or potential species. Anecdotal data from submersible and interactions between deep corals and invasive ROV studies as well as reports from fisherman, species. One invasive species that may be a who have brought them up as bycatch since the threat in the northeast is the colonial tunicate (“sea 19th century, suggests that deep corals have squirt”), Didemnum sp. A. (Bullard et al. 2007). become less common or their distributions have It is currently undergoing a massive population been reduced due to the impacts of bottom explosion and major range expansion and has fishing (Breeze et al. 1997; Watling and Auster become a dominant member of many subtidal 2005). Fishing has had significant effects on deep communities on both coasts of North America. coral populations in other regions. Deep corals It was recently discovered on the northern edge are especially susceptible to damage by fishing of Georges Bank, and is found on hard gravel gear because of their often fragile, complex, substrates, including, pebbles, cobbles, and branching form of growth above the bottom. Also, boulders, and overgrows sessile and quasi- they grow and reproduce at very slow rates, with mobile epifauna (sponges, anemones, scallops, some estimated to be hundreds of years old etc.). The species can occur on immobile sand (Lazier et al. 1999; Andrews et al. 2002; Risk et substrates, but is not known to occur on mobile

211 212 NORTHEAST ofudd eerhr wt is nrdbe rate incredible its with researchers confounded previously has it corals, deep to threat remote a the that is though consensus Even comm.). pers. Valentine, (P. fixed) or the washing of contaminated boat decks or (mobile gear fishing contaminated of use the by or hull theory, boat a in on canyons could, the to brought it be viable, are fragments colonies the because of However, currents. by own or their on far travel cannot they before so hours) settling, few a only for (probably bottom are born from sexual reproduction swim near the that larvae tadpole The margin. bank southern the on located canyons submarine the into area affected present the from getting time difficult a pers. comm.). Valentine, P. comm.; pers. Lambert, (G. higher or >4°C temperatures at be might range a in found preferred its 24°C, is to -2°C of range it temperature though even and 2007), al. et (Bullard Banks Tillies and Stellwagen Georges, only at depths down to 81 m off New England on a shallow water species; so far, it has been found pers. comm.; P. Valentine, pers. comm.). It may be factors in the spread of the tunicate (G. Lambert, temperature, and larval dispersal may be limiting americana Ceriantheopsis Astrangia poculata coral, stony the were overgrow to appear not did that Sound, species epifaunal only Island the Long in bottoms cobble on that pers. comm.) However, Bullard et al. (2007) note MA, Hole, Woods USGS, Valentine, P. comm.; pers. WA, Seattle, Laboratories, Harbor Friday Washington, of University Lambert, (G. feeding did impede least at or tunicate overgrowth, by corals the kill the If Bank. and smother could it habitat, coral deep colonize Georges of Peak Northeast the of substrate gravel the on occur Paragorgia as such corals, those for particularly northeast, to deep coral that occur on hard substrates in the Didemnum (Bullard etal.2007). predation seastar possible of observations some been have there although time, this at predators known species. no the are of There spread rapid and fragmentation of the mats could promote the budding, by rapidly spread to Apotential the has 2007). al. et (Bullard m 45-60 of Bank, at numerous sites over a Georges 230 km On substrate. Didemnum mud or sand STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Didemnum sp. A may be a potential serious threat sp. A covers 50–90% of the substrate and Primnoa , and the cerianthid anemone, Didemnum p wud rbby have probably would A sp. wih r kon to known are which , I adto, depth, addition, In . p i probably is A sp. Didemnum 2 area to a depth Didemnum sp. Asp. sp. monitoring data or published studies from these from studies published or data monitoring affect them. to beany There alsodoesn’tappear to enough deep are U.S. the of coast northeast the along projects coastal the region; of few addition, in this in species coral deep major the to threat significant a pose could that projects etc. mining, mineral cable, pipeline/communication or pending oil and gas exploration/extraction, gas There does not appear to be any specific current Other 2007). throughout substrata al. et (Bullard Canada eastern and England New hard of expanses large colonize eventually could and range its expand and economic damage. The species continues to ecological great cause to potential the has does But comm.). pers. Lambert, (G. time this at spread further its of limits the predict known about this tunicate that it is not possible to is little So expansion. range and colonization of his te atr s prxmtl 40 m to km 400 approximately is latter (the chains Seamount Rise Corner and England New the in seamounts several of surveys were their 2005, and 2004, 2003, In (ROVs). remotely vehicles and operated trawls using England New of off surveys targeted recent, some been have There trends. population long-term any assess clearly to data the using prevent which quantified) or identified properly not were corals (e.g., gaps data many are there but decades, several for bycatch their of part as corals collected have Hatteras Cape to Maine of Gulf the from surveys shellfish and groundfish NEFSC 2004)]. Buhl-Mortensen and (Mortensen Channel Northeast the e.g., waters; coral deep on distribution [There and habitat requirements information in Canadian determine. more to however, is, difficult are trends and severely limited, so their overall population status is requirements, habitat and biology basic their of aspects some as spatial well as U.S., and northeastern temporal the distribution and abundance of deep corals off the surveys, of knowledge faunal our aforementioned the Despite Mapping andResearch VIII. of impactstodeepcorals. types of projects in this region that show evidence RESOURCES AND HABITATS MANAGEMENT OFFISHERY Didemnum sp. A STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION the east of the New England Seamount chain, and nearly midway between the east coast of This action was implemented in May 2005 the U.S. and the Mid-Atlantic Ridge) funded by under Amendment 2 to the Monkfish Fishery NOAA’s Office of Ocean Exploration and National Management Plan (FMP) (NEFMC 2004). EFH Undersea Research Program. The cruises were for some federally-managed species extends multidisciplinary in nature but the goals included beyond the edge of the continental shelf and studying the distribution and abundance of deep includes portions of the canyons. corals relative to the prevailing direction of currents; collecting specimens for studies of reproductive The directed monkfish fishery is conducted with biology, genetics, and ecology; and studying bottom trawls and bottom gillnets, primarily in species associations. Mike Vecchione (NEFSC, coastal and offshore waters of the Gulf of Maine, National Systematics Laboratory) conducted on the northern edge of Georges Bank, and in a multi-year study (2000-2005) exploring the coastal and continental shelf waters of southern faunal biodiversity of Bear Seamount. The multi- New England, including offshore waters on the year program examined some of the temporal edge of the continental shelf and near the heads NORTHEAST variability around the Seamount. Initial results of several of the canyons. Although the current on the biodiversity of deep corals by Moore et al. degree of overlap between the current monkfish (2003) were discussed previously. Mike Fogarty fishing effort and the known presence of corals (NEFSC) in spring 2004 explored the macrofaunal within the canyons is very small, one of the biodiversity of the upper continental slope south fishery management measures contained within of Georges Bank from Oceanographer Canyon Amendment 2, and which was approved by the to Powell Canyon at depths from 400-1100 m, Councils, would increase the probability that the with the deepest stations corresponding to the offshore fishery for monkfish will expand in the shallowest depths sampled on the summit of future. Because there is documented evidence Bear Seamount in the Vecchione survey. of deep corals in the canyons in the area that is identified for possible increased offshore fishing, Fishery Management Councils these closures are intended as a precautionary Fishery management council jurisdiction in the measure to prevent any potential direct or indirect northeast U.S. is primarily the responsibility impacts of an expanded offshore monkfish fishery of the New England Fishery Management on EFH, offshore canyon habitats, and thus, deep Council (NEFMC). In addition to the 26 species corals. under its sole management, the NEFMC shares responsibility over Lophius americanus Approximately 23 federally-managed species (monkfish or goosefish) and Squalus acanthias have been observed or collected within these (spiny dogfish) with the Mid-Atlantic Fishery proposed closure areas, and many of them have Management Council (MAFMC). In 2005, the two EFH defined as “hard substrates” at depths >200 Councils, with the NEFMC as the lead Council on m, which includes habitat or structure-forming the Monkfish Fishery Management Plan (FMP), organisms such as deep corals. Also, the EFH approved the designation of Oceanographer and designations for juvenile and adult life stages of Lydonia Canyons (approximately 116 square six of these managed species (Sebastes spp., nautical miles) as Habitat Closed Areas (HCA) redfish, is one of them) overlap with the two and added these areas to the NEFMC’s network 1 of HCAs (or marine protected areas). These EFH is a provision of the Magnuson-Stevens Fishery Conservation and Management Act (1996). The EFH new HCAs are closed indefinitely to fishing with provision states: “One of the greatest long-term threats bottom trawls and bottom gillnets while on a to the viability of commercial and recreational fisheries monkfish day-at sea (DAS) in order to minimize is the continuing loss of marine, estuarine, and other the impacts of the directed monkfish fishery on aquatic habitats. Habitat considerations should receive Essential Fish Habitat (EFH1) in these deep-sea increased attention for the conservation and man- canyons and on the structure-forming organisms agement of fishery resources of the United States.” therein (Figures 5.1 and 5.8). Within these canyon The definition of EFH in the legislation covers: “those waters and substrate necessary to fish for spawning, habitats, a variety of species have been found breeding, feeding, or growth to maturity.” The legisla- which are known to provide structured habitat, tion mandates that NOAA Fisheries and the Councils including deep corals, and shelter for many implement a process for conserving and protecting species of demersal fish and invertebrates. EFH.

213 214 NORTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF credit: 1999and2001VTR). (1998) andWatling etal.(2003)databases; and,1999and2001directedmonkfishottertrawltrips. Source 2 totheMonkfishFishery Management Plan;locationsofknownalcyonaceans fromthe Figure 5.8. The OceanographerandLydonia CanyonEssentialFishHabitatclosedareas, under Amendment Theroux andWigley STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION closed areas, and EFH for all six of these species document the presence of corals in the other 10 has been determined to be vulnerable to bottom canyons. trawling and perhaps also vulnerable to bottom gillnets. Although deep corals are not explicitly The New England Fishery Management Council included in the EFH descriptions for any species alone has also indefinitely closed an additional in the northeast region, some deep corals are, 3,000+ square nautical miles, as Habitat Closed of course, known to grow on hard substrates, Areas, in the Gulf of Maine, Georges Bank, and and may themselves be considered a form of southern New England to bottom-tending mobile substrate. The rationale is that, since there are fishing gear to protect EFH (Figure 5.1), which corals found within these proposed closed areas, indirectly protects any deep corals in those this is indicative of areas of hard bottom. Also some areas. This initial suite of HCAs was created coral species may provide the structural attributes under both Amendment 10 to the Atlantic Sea of habitat similar to that provided by other dense Scallop Fishery Management Plan in 2003 and epifaunal assemblages (as discussed above), Amendment 13 to the Northeast Multispecies and may be particularly vulnerable to damage Fishery Management Plan in 2004 as “Level 3” NORTHEAST or loss by trawling or gillnets. Thus, by avoiding closures, and are closed indefinitely to all bottom- any direct adverse impacts of bottom trawls and tending mobile gear to protect EFH. The canyon gillnets used in the monkfish fishery on EFH for habitat closures implemented under Amendment the six species of fish and any indirect adverse 2 to the Monkfish FMP should be viewed as an impacts on hard bottom habitat and emergent addition to the suite of HCAs, and in concert epifauna, such as the deep corals, that grow in with the other HCAs, provides a large network of these habitats within the closed areas, adverse marine protected areas (MPAs) as compared to impacts of an expanded offshore fishery would the relatively small size of the geographic region be minimized to the extent practicable. under management.

Protection of deep corals is a relatively new Lastly, under the current effort to update all of the concept in this region and the NEFMC believes EFH provisions of all of the NEFMC FMPs, the that there are several statutory and regulatory NEFMC has approved a set of new Habitat Areas authorities that support the Councils’ initiative to of Particular Concern (HAPCs2) including many protect deep coral habitats. The NEFMC took this steep-walled canyons on the eastern seaboard proactive and precautionary approach to protect extending from the Hague Line to the North these sensitive habitats through aggressive Carolina/South Carolina border and Bear and fishery management measures, which is based Retriever Seamounts to protect sensitive EFH on sound ecological principles, as appropriate and the habitat and structure-forming organisms and necessary. In Amendment 2 to the Monkfish therein. These new HAPCs will need approval FMP, the Councils, for the same reasons and from the MAFMC before they can be implemented rationale discussed above, also considered a through final action sometime in late 2008. The management alternative that would have closed MAFMC will likely take up the topic at one of their all 12 steep-walled canyons along the continental upcoming meetings shelf-break from the Hague Line in the north to the North Carolina/South Carolina border in the IX. REGIONAL PRIORITIES TO south. Although this management alternative UNDERSTAND AND CONSERVE DEEP was not ultimately chosen by the Councils CORAL COMMUNITIES for implementation due to the lack of readily available coral data and potential negative social Mapping and economic impacts, the Councils did feel that Deep corals have been largely unmapped off the the science and data supported the closure of northeastern U.S, particularly in the Mid-Atlantic. Oceanographer and Lydonia Canyons at that What is currently known about coral distribution time as a precautionary measure. The Councils 2 determined that protection of deep corals in all 12 HAPCs are a subset of the much larger area identified as EFH that play a particularly important ecological canyons would be less certain than in just closing role in the life cycle of a managed species or that are Lydonia and Oceanographer Canyons, until rare and/or particularly sensitive or vulnerable to such time as additional surveys are conducted human-induced environmental degradation and or evidence is examined which more thoroughly development activities.

215 216 NORTHEAST no oa lra dsesl Gntc tde may studies Genetic dispersal. larval coral into used, and this line of research may provide insight be could that tool one is genetics Molecular out. their some basic taxonomic issues need to be worked and difficult, step, first a as so question, in is often is taxonomy corals deep of Identifying identifications the surveys. historical various the from of corals deep accuracy the about uncertainty is there that previously stated was It Research deep toward coral conservation. step important an be would U.S. suitable habitat for deep coral in the northeastern to generate habitat suitability maps and to identify modeling predictive of use The 5-10°C. between speed, rocky substrates and a temperature range distributed and located in areas with high current sloped for steeply habitat in the while substrates, rocky on and environments predominantly occurred requirements habitat gorgonians. two the between differed the that showed Their analysis. results the in included were substrate break. chlorophyll shelf temperature, slope, including and factors environmental shelf Several continental Atlantic resedaeformis (2005) Primnoa for habitat Metaxas suitable of and areas determine to models predictive developed Leverette Recently, should beestablished. programs be monitoring long-term to and need assessed, abundance and distribution and coral (McDonough exist to Puglise 2003). Temporal/spatial changes in deep known are corals deep where areas site-specific for produced should be maps high-resolution and corals, deep of locations potential identifying of purposes for areas large cover that produced be should maps Low-resolution etc. sleds, camera towed ROVs, and multibeam sonars, manned submersibles and side-scan as such technologies advanced more their habitats, particularly in the canyons, utilizing to identify, map, and characterize deep corals and for management purposes. Therefore, it is critical data temporally to provide distributional of data adequate density and the spatially diffuse too much is generated cases they most in vast, surveys such was of breadth the While ago. years more or twenty done was that cameras drop and samplers, grab gear, trawl with sampling grid or random, blind, on based largely is region this in REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF . resedaeformis P. aaoga arborea Paragorgia a ln te Canadian the along cret pe and speed current , a mr broadly more was P.arborea and

eerh n t hr crl aooit at taxonomists coral hire museums anduniversities. to taxonomic for and available research be to needs funding more and level, graduate the at taxonomy coral for example. More students need to be trained in make to difficult be distribution, and mapping coral deep in progress will it taxonomists, professional few coral so With corals. deep identify properly to available taxonomists coral qualified of shortage a is there that note to important it’s and Maine of Gulf However,offBank. canyons Georges submarine deep the into Seamounts England New the from dispersing are corals the whether e.g., distributions; geographic broader with species of populations simply or endemic, are Seamounts England New the around corals the whether determine to is used being technology currently Atlantic. DNA-sequencing the example, of side For either on and regionally,nationally, corals comparing for useful be also yac qatt i te NEFSC the magellanicus in quantity bycatch observer fisheries and program logs. Prior to the surveys year 2000, for example, shellfish and groundfish the in bycatch coral deep their about quantitative more become to needs NEFSC The Brock (2003). in found be can McDonough and Puglise (2003) and Puglise and corals deep on research for recommendations these of Many destroyed. or rates of injured been have the recovery that habitats coral and corals to quantify and fishing, from especially threats, anthropogenic various to corals deep of resilience or vulnerability quantify the to necessary is it Finally, determined. as EFH for Federally managed species has to be terms of the latter, In the possible quantified. role of and deep corals described be should species associated of histories life the in play corals the relationships need to be defined, and the role that coral Deep habitat habitats. biodiversity should be coral assessed, food deep web the of extent understand and distribution the to affect that order parameters habitat physical in other data and parameter associated geologic, collect In to oceanographic, characterized. important is be it addition, to need requirements coral habitat Their feeding. and deep rates, recruitment, recolonization on reproduction, physiology, questions growth, fundamental are There region. this in needed are corals deep on taxonomy,to studies addition history In life basic Alni sa clo) surveys scallop) sea (Atlantic Placopecten STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION were estimated by cursory visual inspection an increased mapping and survey effort at the or “eyeballing” only (D. Hart, NOAA Fisheries Federal and academic level (including joint Service, NEFSC, Woods Hole Laboratory, Woods studies with Canada); 2) more basic research Hole, MA, pers. comm.). The bycatch data for on deep coral taxonomy, life history, habitat those surveys were divided up into 3 categories: requirements, species associations, etc.; and substrate, shell, and other invertebrates; and the finally, 3) quantification on the susceptibility log sheets only recorded percent composition and of deep corals to anthropogenic influences, total volume (bushels). In the fisheries observer particularly fishing. program, the observers also log the presence of coral bycatch; however, they are lumped into one category (“corals and sponges”), and are not identified further. In addition, because the observer program observes thousands of trips every year in dozens of different fisheries, with each fishery having its own regulations for mesh NORTHEAST size and configuration, a reported absence of coral at a location may simply be a function of the catchability of the gear used (D. Potter, Fisheries Sampling Branch, NOAA Fisheries Service, NEFSC, Woods Hole Laboratory, Woods Hole, MA, pers. comm.). This is also a problem with the NEFSC surveys; it is important to remember that fishing gear is not designed to “catch” corals. But at the very least, if there was an attempt made to properly identify coral bycatch from these two programs, one might come up with some form of presence data.

X. CONCLUSION

The overall quantity of deep coral habitat in the northeast region is unknown, and no systematic assessment of the distribution, abundance, and population dynamics of deep coral is available for this region. That, along with a dearth of information on their natural history, as well difficulties with their taxonomy, makes it difficult, if not impossible, to determine if there have been changes in deep coral occurrence or abundance over time. Nevertheless, even though there is no quantitative information on the extent of anthropogenic impacts to deep corals in this region, some of the areas where structure-forming deep corals are definitely known to occur (e.g., unique areas such as the submarine canyons and the New England Seamounts) are currently protected or under consideration for protection from bottom-tending fishing gear as EFH and Habitat Areas of Particular Concern.

Obviously, in order to better preserve and protect deep corals and deep coral habitat off the northeastern U.S., there needs to be 1)

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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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221 222 NORTHEAST Watling L (2005) Deep-sea octocorals as homes Verrill AE (1922) The Alcyonaria of the Canadian remarkable the of Notice (1884) AE Verril to additions recent of Notice (1879) AE Verrill to additions recent of Notice (1878b) VerrillAE to additions recent of Notice (1878a) VerrillAE a of Notice (1862) AE Verrill (1980) RA Cooper JR, Uzmann PC, Valentine In: geology. Submarine (1987) BE Tucholke the of overview An (1992) DW Townsend REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Jones andBartlettPublishers.pp56-113 MA: Boston, rise. and slope The continental (eds) Atlantic U.S. the of WR environment marine Wright, JD, Milliman, Ocean Coastal NOAAProg. Reg.Synth.Ser. 1 Maine. of The Gulf (eds) In: PF 5-26 Larsen, Pages: DW, Maine. Townsend, of Gulf the of productivity biological and oceanography commensals.html> 04mountains/background/commensals/ oceanexplorer.noaa.gov/explorations/ and the Sea 2004/Coral Commensals. < Explorer/ Oceanic in Administration/Mountains Atmospheric Ocean National species. Explorations. other for VIII: etc. PartG: coelenterates, Alcyonaria and Actinaria vol echinoderms, 1913-18, molluscs, Expedition Arctic Canadian the from Reports species. and a with genera Canadian other some of revision 1913-1918, Expedition, Arctic Arts and Series 3,28:213-2 Science of Journal England. American New of coast southern the of banks outer the occupying fauna marine of Scienceand Arts Series3,17:472-474 Journal American 5. No. America, North of coast eastern the of fauna marine the of Scienceand Arts Series3,16:371-379 Journal American 2. No. America, North of coast eastern the of fauna marine the of Science and Arts Series3,16:207-215 Journal American America. coast North eastern of the of fauna marine the MA, USA,3:127-129 Salem, Inst., Essex Proc. Bank. Georges 283-312 38: Geology Marine canyon. submarine Oceanographer of biology and Geology

Primnoa from http:// Watling L, Moore J, Auster P (2005) Mapping the Hecker C, Skinder I, P,Babb Auster L, Watling deep- of Distribution (2005) P L, WatlingAuster Wigley RL (1968) Benthic invertebrates of the of invertebrates Benthic (1968) RL Wigley Caron RH, Backus EH, Backus PH, Wiebe Watling L, Mosher C (2006) Commensalism in (2006) C Mosher A, Simpson L, Watling onan i te e 2004/Mission < gov/explorations/04mountains/logs/ Sea the Summary. in Mountains National Oceanic and Atmospheric Administration/ Explorer/Explorations. seamounts. of Ocean biodiversity overall the assessing and octocorals of distribution Connecticut, Groton,USA National Center,Research Undersea of University the CD-ROM. 1.0 Version of slope. database and shelf continental alcyonaceans geographic U.S. northeastern A deepwater (2003) of B Berlin, Heidelberg Springer-Verlag ecosystems. and corals Freiwald A, in Roberts JM (eds.), 259-264 Cold-water Pages States. United the of coast northeast the off water Alcyonacea Naturalist 5:8-13 Underwater banks. fishing England New Jones andBartlettPublishers MA: Boston, rise. The and slope continental (eds.) Atlantic U.S. the WR In: of environment marine Wright 140-201. JD, Biological Milliman (1987) Pages oceanography. JB Powers Waterbury JF, Grassle K, PM., Gilbert DA, html 05stepstones/logs/summary/summary. < and oceanexplorer.noaa.gov/explorations/ Administration. Oceanic Atmospheric National Stepping Stones/Log. Atlantic Explorations/North Explorer/ Ocean hosts. octocoral their to the deep sea: relationships of invertebrates logs/summary/summary.html < Atmospheric noaa.gov/explorations/05stepstones/ and Administration. Stones/Log. Oceanic Stepping National Atlantic North Explorer/Explorations/ Ocean octocorals. deep-sea of distribution and Taxonomy summary/summary.html > http://oceanexplorer.noaa. http://oceanexplorer. > > http:// STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION Reference Cairns and Chapman 2001; Moore et al. 2003 Cairns and Chapman 2001 Hecker 1980; et al. 1983; Hecker 1990; Cairns and Chapman 2001; Guida (unpublished data) Cairns and Chapman 2001 Hecker 1980; and Blechschmidt 1980; Hecker et al. 1980, 1983; et al. 1982; Malahoff Cairns and Chapman 2001; Moore et al. 2003 NORTHEAST 37-366 183-1646 403-2634 183-2250 1487-2286 Depth Range (m) Distribution Widespread (cosmopolitan) distribution; found on Bear Seamount Atlantic Endemic to western Widespread (cosmopolitan) distribution. Continental slope south of New England, Canyon, continental shelf between Lydonia Baltimore and Hudson Canyons, in Canyon, and between 100-200 m on the shelf south of Hudson Canyon and in the head Hudson Canyon Amphi-Atlantic with a disjunct distribution Widespread (cosmopolitan) distribution; found Heezen, Lydonia, in several canyons (Corsair, Baltimore, Norfolk; near Oceanographer, Hudson); continental slope on the southwestern and edge of Georges Bank, between Veatch Hydrographer Canyons; in the Mid-Atlantic on the slope between Lindenkohl Canyon on south and Carteret Canyon on the north; in Toms Mid-Atlantic on the slope bounded by Canyon to the south and Meys north; Bear Seamount

Cairns, 1979 Alcock, 1898 Species Caryophyllia ambrosia ambrosia Caryophyllia ambrosia caribbeana Dasmosmilia lymani (Pourtales, 1871) Deltocyathus italicus (Michelotti, 1838) Desmophyllum dianthus 1794) (Esper, List of deep coral species found in the waters off the Northeastern United States. ** = distribution information based on studies or surveys of a List of deep coral species found in the waters off Higher Taxon Phylum Cnidaria Anthozoa Class Subclass Hexacorallia Order Scleractinia Family Caryophillidae Appendix 5.1. particular area of the Northeast Region, not on overall distribution.

223 224 NORTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Higher Taxon Species Distribution Depth Range (m) Reference Lophelia pertusa (L, 1758) Widespread (cosmopolitan) distribution; 146-1200; 700-1300 Hecker 1980; Hecker Oceanographer Canyon wall; Bear Seamount and Blechschmidt 1980; Hecker et al. 1980; Cairns and Chapman 2001; Moore et al. 2003 Solenosmilia variabilis Widespread (cosmopolitan) distribution; Lydonia 220-1383 Hecker 1980; Hecker Duncan, 1873 canyon; on the slope bounded by Toms Canyon et al. 1983; Cairns and to the south and Meys Canyon to the north Chapman 2001

Vaughanella margaritata Endemic to northwestern Atlantic; Bear 1267 Cairns and Chapman (Jourdan, 1895) Seamount 2001; Moore et al. 2003 Family Dendrophylliidae Enallopsammia profunda Endemic to western Atlantic 403-1748 Cairns and Chapman 2001 (Pourtales, 1867) Enallopsammia rostrata Widespread (cosmopolitan) distribution 300-1646 Cairns and Chapman 2001 (Pourtales, 1878) Family Flabellidae Flabellum alabastrum Amphi-Atlantic with contiguous distribution; 357-1977 Hecker 1980; Hecker Moseley, 1873 Canyons (Corsair, Heezen, Lydonia and Blechschmidt 1980; Oceanographer, Alvin, Baltimore, Norfolk) Hecker et al. 1980, 1983; and slopes; Bear Seamount. some may be F. Cairns and Chapman angularis or F. moseleyi 2001; Moore et al. 2003

Flabellum angulare Amphi-Atlantic with contiguous distribution; see 2266-3186 Hecker 1980; Hecker Moseley, 1876 also F. alabastrum and Blechschmidt 1980; Hecker et al. 1980, 1983; Cairns and Chapman 2001; Moore et al. 2003 Flabellum macandrewi Amphi-Atlantic with contiguous distribution; see 180-667 Hecker 1980; Hecker Gray, 1849 also F. alabastrum and Blechschmidt 1980; Hecker et al. 1980, 1983; Cairns and Chapman 2001; Moore et al. 2003 Javania cailleti (Duch. & Widespread (cosmopolitan) distribution; 30-1809 Hecker 1980; Hecker Mich., 1864) Lydonia, Oceanographer Canyons et al. 1983; Cairns and Chapman 2001 Family Fungiacyathidae Fungiacyathus fragilis Widespread (cosmopolitan) distribution 412-460 Cairns and Chapman 2001 Sars, 1872 STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION Reference Theroux and Wigley 1998; Cairns and Chapman 2001 Brugler 2005 Smithsonian Institution collections Auster 2005 and Watling Hecker and Blechschmidt 1980; Hecker et al. 1980, 1983; Opresko 1980; et al. 1980; Valentine Hecker 1990; Moore et al. Auster and 2003; Watling 2005 Hecker and Blechschmidt 1980; Hecker et al. Opresko 1980; Watling Auster 2005 and Auster 2005 and Watling Hecker and Blechschmidt 1980; Hecker et al. Opresko 1980; Watling Auster 2005 and NORTHEAST 262 0-263 750-1326 700-2600 750-1099 1643, 1754 Depth Range (m) Distribution Endemic to western Atlantic Endemic to western Near and on Bear Seamount Virginia Off Hard substrates from Corsair Canyon to Hudson Canyon; outcrops in Corsair Oceanographer Canyons; in Heezen, Lydonia, Alvin Canyon; on slope the on slope near southwestern edge of Georges Bank, between and Hydrographer Canyons; in Mid- Veatch Atlantic on slope flanked by Lindenkohl Canyon to south and Carteret Canyon north on Canyon to south and Toms slope bounded by Meys Canyon to north; Bear Seamount Soft substrates, highest densities in canyons; Heezen, Oceanographer found in Corsair, Toms Canyons; seen near Hudson Canyon, Canyon, in Baltimore axis of Norfolk Canyon Found in axis of Heezen, Lydonia, Oceanographer Canyons

(Ellis & (Verrill, (Verrill, sp.** sp.** Species Astrangia poculata 1786) Solander, Leiopathes Cirrhipathes Alcyonium digitatum Linné, 1758 Anthomastus agassizii 1922 ** Verrill, Anthomastus grandiflorus 1878 ** Verrill, Clavularia modesta 1874) (Verrill, Clavularia rudis 1922)** Higher Taxon Family Rhizangiidae Antipatharia Order Antipathidae Family Subclass Octocorallia Alcyonacea Order Alcyoniidae Family Family Clavulariidae

225 226 NORTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Higher Taxon Species Distribution Depth Range (m) Reference Family Nephtheidae Capnella florida (Rathke, Lydonia, Oceanographer, Baltimore Canyons; 350-1500 Hecker and Blechschmidt 1806)** axis of Heezon Canyon; wall of Corsair Canyon; 1980; Hecker et al. 1980; continental slope south of New England off Opresko 1980; Hecker Georges Bank 1990; Watling and Auster 2005 Capnella glomerata Several individuals found in Lydonia Canyon 200-561 Hecker et al. 1980; (Verrill, 1869)** Opresko 1980; Watling and Auster 2005 Gersemia fruticosa** Near and in deep portion of Hudson Canyon; at 600-3100 Hecker and Blechschmidt (Sars, 1860) the mouth of Norfolk Canyon; seen near heads 1980; Opresko 1980; of Toms and Carteret Canyons (i.e., between Watling and Auster 2005 Baltimore and Hudson Canyons)

Gersemia rubriformis Watling and Auster 2005 (Ehrenberg, 1934) Order Gorgonacea Family Acanthogorgiidae Acanthogorgia armata Found in many canyons (Corsair, Lydonia, 350-1300 Hecker and Blechschmidt Verrill, 1878 ** Oceanographer, Alvin, near Hudson, Norfolk, 1980; Hecker et al. 1980; Baltimore); seen on boulders or outcrops Opresko 1980; Malahoff et al. 1982; Watling and Auster 2005 Family Anthothelidae Anthothela grandiflora Lydonia, Oceanographer, Baltimore Canyons 450-1150 Hecker et al. 1980; (Sars, 1856) ** Opresko 1980; Watling and Auster 2005 Family Chrysogorgiidae Chrysogorgia agassizii Several individuals that may be C. agassizii 2150 Watling and Auster 2005 (Verrill, 1883) found in the vicinity of Hudson Canyon.

Iridogorgia pourtalesii Watling and Auster 2005 Verrill, 1883 Radicipes gracilis (Verrill, Watling and Auster 2005 1884) STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION Reference Hecker and Blechschmidt 1980; Hecker et al.1980; Opresko 1980; Hecker Theroux and Wigley 1990; Auster and 1998; Watling 2005 Auster 2005 and Watling Auster 2005 and Watling Moore et al. 2003; Watling Auster 2005 and Wigley 1968; Hecker and Blechschmidt 1980; Hecker et al. 1980; Theroux Opresko 1980; and Grosslein 1987; Theroux and Wigley 1998; Moore et al. 2003; Watling Auster 2005 and Hecker and Blechschmidt 1980; Hecker et al. 1980, 1983; Opresko 1980; et al. 1980; Valentine Auster 2005 and Watling NORTHEAST 300-1100 600-2000 400-2200 Depth Range (m)

Distribution Found in Corsair, Heezen, Oceanographer Found in Corsair, Alvin, Baltimore Canyons; on slope near Canyons; in Mid-Atlantic on slope flanked by Lindenkohl Canyon to south and Carteret Toms Canyon to north and on slope bounded by Canyon to south and Meys north; continental slope south of New England off Georges Bank; seen on soft substrates Bear Seamount? Found in Gulf of Maine, Georges Bank, and Baltimore, Oceanographer, Canyons (Lydonia, Norfolk); probably Bear Seamount Found in Gulf of Maine and canyons from Corsair to near Hudson, seen in Corsair, Canyons; Lydonia Heezen, Oceanographer, Alvin Canyon; on slope the on slope near southwestern edge of Georges Bank, between and Hydrographer Canyons; in Mid- Veatch Atlantic on slope flanked by Lindenkohl Canyon to south and Carteret Canyon north on Canyon to south and Toms slope bounded by Meys Canyon to north

Verrill, Verrill, Wright, Wright, grandis

grayi ornata

caryophyllia arbuscula Species

Acanella (Johnson, 1862) ** Keratoisis 1869 Keratoisis 1878 Lepidisis 1883** Verrill, Paragorgia arborea (Linné, 1758) ** Paramuricea 1883 ** Verrill, Higher Taxon Family Isididae Family Paragorgiidae Family Plexauridae

227 228 NORTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Higher Taxon Species Distribution Depth Range (m) Reference Paramuricea placomus Gulf of Maine Watling and Auster 2005 (Linné, 1758) ** Paramuricea n. sp. Watling and Auster 2005 Swiftia casta (Verrill, Bear Seamount? Moore et al. 2003; Watling 1883)** and Auster 2005 Family Primnoidae Narella laxa Deichmann, Watling and Auster 2005 1936 Primnoa resedaeformis Found in Gulf of Maine, Georges Bank, and 91-548 Hecker and Blechschmidt Gunnerus, 1763) ** Canyons (Lydonia, Oceanographer, Baltimore, 1980; Hecker et al. 1980, Norfolk); south to off Virginia Beach, VA; 1983; Opresko 1980; probably Bear Seamount Valentine et al. 1980; Theroux and Grosslein 1987; Theroux and Wigley 1998; Moore et al. 2003; Cairns and Bayer 2005; Watling and Auster 2005; Heikoop et al. 2002. Thouarella grasshoffi Watling and Auster 2005 Cairns, 2006 Order Pennatulacea Family Anthoptilidae Anthoptilum grandiflorum Newfoundland to Bahamas, Lousiana, Chile, 274-3651 US NMNH collection, Hawaii, Antarctica, N. Europe OBIS Anthoptilum murrayi Lydonia Canyon to Puerto Rico, Hawaii, 430-2491 (1538 m US NMNH collection, Aleutians, Japan, W. Africa, N. Europe min in NE US) OBIS Family Kophobelemnidae Kophobelemnon Newfoundland to South Carolina, Japan, W. 393-2199 (1330 m US NMNH collection, stelliferum Africa, N. Europe min in NE US) OBIS Kophobelemnon scabrum Nova Scotia to Virginia 1977-2249 US NMNH collection Kophobelemnon tenue Massachusetts to Virginia 2491-4332 US NMNH collection Family Pennatulidae Pennatula aculeata Newfoundland to Virginia, California, Iberia, N. 119-3316 US NMNH collection, Africa OBIS Pennatula grandis New Jersey, Bay of Biscay 1850-2140 US NMNH collection, OBIS Pennatula borealis Newfoundland to North Carolina, California 219-2295 US NMNH collection, OBIS STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION Reference US NMNH collection, OBIS US NMNH collection US NMNH collection, OBIS US NMNH collection US NMNH collection, OBIS US NMNH collection US NMNH collection, OBIS US NMNH collection US NMNH collection, OBIS NORTHEAST NE US) NE US) 1483-2359 1334-2194 2513-4332 1502-2505 min in NE US) min in NE US) report at 59 m) Depth Range (m) 549-3338 (1538 m 2683-3740 (3166 m 1211-2844 (doubtful 1211-2844 20-812 (51 m min in 37-2249 (229 m min in Distribution Nova Scotia to North Carolina, W. Africa, N. Nova Scotia to North Carolina, W. Eurpoe Nova Scotia to Virginia Africa, N. Massachusetts to North Carolina, W. Europe Massachusetts to Virginia Massachusetts to North Carolina, Panama, W. Africa Louisiana Massachusetts to Virginia, Islands, Louisiana, Massachusetts to the Virgin Suriname, N. Europe, Indian O. Newfoundland to Massachusetts, NC (doubtful), Alaska Islands, the Virgin Iberia York-Florida, New Species Distichoptilum gracile Protoptilum abberans Protoptilum carpenteri Scleroptilum gracile Scleroptilum grandiflorum Umbellula guntheri Umbellula lindahlii Balticina finmarchica Stylatula elegans Higher Taxon Family Protoptilidae Family Scleroptilidae Family Umbellulidae Family Virgulariidae ** Distribution information based on studies of a particular area in the Northeast Region, not an overall distribution.

229 230 NORTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF Appendix 5.2. Deep coral species discussed in Hecker and Blechschmidt (1980), Opresko (1980) (octocorals), and Hecker (1980) (scleractinians), as well as Hecker et al. (1980). Species names are listed exactly as stated in the literature. SPECIES DISTRIBUTION OR LOCATION SCLERACTINIANS: Stony corals Continental shelf between Baltimore and Hudson Canyons, in Baltimore Canyon, and between 100-200 m on the shelf south Dasmosmilia lymani of Hudson Canyon and in the head of Hudson Canyon; soft substrates. Same as D. dianthus. Outcrops and underhangs at depths from 1000-1900 m. Seen on outcrops in Corsair Canyon. Found in Heezen Canyon. Seen in deeper parts of Lydonia Canyon, and on boulders or outcrops in Oceanographer Canyon, between Desmophyllum cristagalli 650-1600 m. Found on an outcrop near Hudson Canyon. Occasionally in axis of Norfolk Canyon. Canyons and slope from 600-2500 m; some may be F. angulare or F. moseleyi. Seen in Corsair Canyon. Found in Heezen and Oceanographer Canyons on soft substrate. Seen on deep continental slope near Alvin Canyon. Found on slope south of Baltimore Canyon. Found in deeper parts of the continental slope south of Norfolk Canyon and in axis of Norfolk Canyon on Flabellum alabastrum soft substrate. Same as L. pertusa. West wall of Oceanographer Canyon at 1100 m; dead rubble also found on wall at depths from 700-1300 Lophelia prolifera m. Solenosmilia variabilis Large colony recovered from the east flank of Lydonia Canyon. Javania cailleti One specimen recovered in axis of Oceanographer Canyon between 935-1220 m. ALCYONACEANS: Soft corals Soft substrates, highest densities in canyons. In the northern canyons found from 700-1500 m, southern canyons from 1500- 2200 m; as deep as 2600 m. Found in Corsair, Heezen (west wall), and Oceanographer Canyons. Seen near Hudson Canyon, Toms Canyon, in Baltimore Canyon, and in axis of Norfolk Canyon. Frequently seen where a species of Pennatula was also Anthomastus grandiflorus common. Hard substrates from Corsair Canyon to Hudson Canyon from 750-1900 m. Seen on outcrops in Corsair Canyon. Found in Heezen Canyon. Seen in deeper parts of Lydonia Canyon. On boulders or outcrops in Oceanographer Canyon; 1057-1326 m. Seen on deep continental slope near Alvin Canyon. Seen near heads of Toms and Carteret Canyons (i.e., between Baltimore Anthomastus agassizii and Hudson Canyons). Same as Gersemia fructicosa (?). Southern part of study area at depths from 2300-3100 m. Seen near Hudson Canyon around 2250-2500 m and at the mouth of Norfolk Canyon; populations found in deep portion of Hudson Canyon. Seen near heads of Toms and Carteret Canyons (i.e., between Baltimore and Hudson Canyons). Different form seen in Corsair and Heezen Eunephthya fruticosa Canyons between 600-1200 m may be E. florida (see Opresko 1980) (Same as Capnella florida?) Same as Capnella florida (?). Found in Lydonia, Oceanographer, Baltimore Canyons, but only high abundances in Lydonia at Eunephthya florida 350-1500 m. Axis of Heezon Canyon between 1100-1200 m; wall of Corsair Canyon between 600-1000 m. Eunephthya glomerata Same as Capnella glomerata (?). Several individuals found in Lydonia Canyon at 200 m and 562 m depth. Same as Clavularia rudis. Axis of Heezen Canyon at 1100 m, Lydonia Canyon at 900 m, Oceanographer Canyon at 750 m and Trachythela rudis 900 m. STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION NORTHEAST Found from Corsair Canyon to a site near Hudson Canyon at depths of 700- of depths at Canyon Hudson near site a to Canyon Corsair from Found . (?) were found at 2150 m in the vicinity of Hudson Canyon. placomus . agassizii P . C . Found in Lydonia Canyon at 560 m, in Baltimore Canyon at 450 m, and Norfolk Canyon at 400 m. 400 Canyon at Norfolk and m, 450 Canyonat Baltimore in m, 560 at Canyon Lydonia in Found . , perhaps also perhaps , resedaeformis grandis . . P P DISTRIBUTION OR LOCATION Norfolk m, 500 and m 400 Canyon Baltimore of axis m, 300-1100 around Canyon Oceanographer m, 300-900 Canyon Lydonia Canyon 400-600. m. and Baltimore Canyons between 450-1149 Oceanographer, Found in Lydonia, found Canyons; Oceanographer and Corsair in outcrops or boulders on Seen depth. m 600-2500 from canyons many in Found and Oceanographer Alvin Canyons Canyon. between Found 400-1299 on in m. Seen Lydonia on deep continental slope near an outcrop near Hudson Canyon. Found at 350 m in Baltimore Canyon. Occasionally in axis of Norfolk Canyon on exposed outcrops. Found from Corsair Canyon to Hudson Canyon between 750-2150 m. Found on and Oceanographer Canyons. found at depths between 400-1349 m in Lydonia wall and axis of Oceanographer Canyon; as Same 2200 m on hard substrates. Seen on outcrops in Corsair Canyon. Found in Heezen Canyon. Seen in deeper parts of Lydonia Canyon. On boulders or outcrops in Alvin Oceanographer Canyon. Not Canyon. seen Seen in on deep continental slope near Norfolk Canyon. as Same On soft substrates from 600-1300 m depth in the Alvin north near slope and continental 1500-2000 deep on m Seen m. depth in 1046-1191 between the Canyon Oceanographer south. in Seen Found Canyons. in Oceanographer Corsair, Heezen, and species. Canyon. On slope just south of Baltimore Northern and southern forms may be different Several individuals that may be

SPECIES GORGONACEANS: Gorgonians Paragorgia arborea Anthothela grandiflora Acanthogorgia armata Paramuricea grandis Paramuricea borealis Primnoa reseda Acanella arbuscula Chrysogorgia agassizii

231 232 NORTHEAST STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION US NORTHEASTERN THE IN ECOSYSTEMS STATE CORAL DEEP OF