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IDENTIFICATION, DISTRIBUTION, AND CONSERVATION OF DEEP-SEA CORALS IN

CANADA'S NORTHWEST ATLANTIC

by

in partial fulfillment of therequirementsfor the degree of

Memorial University April 2010

ABSTRACT

Deep-sea corals are long-lived,slow-growing benthic animals and are generally considered important for deep-sea biodiversity.Deep-seacorals in Newfoundland, Labrador,and eastem Canadian Arctic waters were mapped using incidentalby-catch from multispecies scientific surveys and fisheries observations.Todate(2004-2009),44 deep-sea coral species have beendocumented,including 330ctocorals,eight scleractiniansandthreeantipatharians.Coraldistributionswerehighlyclustered,with most species co-occurring in fishing sets.Fivecoral species diversityandabundance hotspotswere delineated:HudsonStrait regioin,LabradorsheIf edge and slope,Orphan Spur-Tobin'sPoint,Flemish Passand sounlwestGrandBanksshelfedgeandslope Corals are under threatfrombottomtendingfishing.Impactsfrom mobile andfixed gears caninclude dislodgement,breakage,and completeremoval. Althoughseveral protected areashavebeenestablishedand other candidateshave been identified, protective measures for deep-seacoralin Newfoundland andLabradorareinsufficient

ACKNOWLEDGEMENTS

Iwishto thank Dr.Evan EdingerandDr. RichardHaedrichfromMemorial Universityof Newfoundland(MUN), andDr. KentGilkinson andJasonSimms fromDepartmentof Fisheriesand Oceans Canada (DFO) for their support and guidance throughout this project.Iwould like to thank all theNewfoundlandandLabrador,fisheriesobservers (SeaWatchLtd.,St.John's,NL),techniciansandscientificstaff(DFO,MUN),sea-going personnel from the Canadian Coasl Guard Ship (CCGS) Teleost,GGGS Templeman, GGGSHudson,andResearchVessel(RV) GapeBallardfortheir cooperation and contributions to the project.Thankstolocalfishers fortheirinsight.Specialthanksto

moralsupport.Thanks to Dr.R.Devillers(MUN),T.Bowdring(DFO),D.Reddick(DFO), G.Cossett(DFO),andN.Ollerhead(DFO)fortheirassistanceandguidancein generatingmapsandposters.Susan Gass providedguidance onmethodologiesand exploratorydata from earlier researchstudies.DepartmentofFisheriesandOceans Science division provided logisticalsupport.This projectwas supportedbytheDFO OceansDivisionandInternational Goverance Program, NationalScience and Engineering ResearchCouncil (NSERC)DiscoveryGrant toDr. E.Edinger,and by MUN.Thanks to Rose and PatVeitch for editing.Finally,special thankstomy family (partnerMike,myparentsBrettandEvelyn,andmybrotherBrettF.andfamily)and exceptionalfriends(Angie,Tanya,Susanna, and Jen) for their ongoing support, never- endingpatience,andmostofallfortheirfaithforbelievinginme

Dedicatedto my AuntElaine(1947-199S), myrolemodel and inspiration in life

TABLE OF CONTENTS

1 Introduction to Deep-Sea CoralInThe Newfoundland and LabradorReglon,

1.1.6 Where Do Deep-Sea Corals Live?

1.1.7Ecological Importance

1.2 ThreatstoDeep-SeaCorals 1.2.1 Bottom Fishing Practices

Hydrocarbon Exploration

ResearchSurveys Climate Change

2 DistributionofDeep-Sea Coralsin theNewfoundlandandLabradorRegion,

2.3.1MultispeciesStock Assessment Surveys andthe NorthernShrimp

2.3.2Fisheries Observer Program

2.3.3DefinitionofCoralSpeciesRichness Hotspotsand Abundanee Peaks

2.3.4MappingofDeep-seaCoral

2.4.1Multispecies Stock AssessmentSurveys andNorthernShrimp Survey

2.4.3Deep-SeaCoralDistributionandDiversity Patterns

2.5.2OtherAreas ofInterest(AOI)

2.5.3 Substrates of CoralBiodiversityHotspots and Other Areasoflnterest

2.5.4 Comparison withLocal Ecological Knowledge

2.5.6Limitationsand ConservationImplications

3 UpdatesonDeep-SeaCoraIDistributionintheNewfoundland,Labrador,and ArcticRegions, NorthwestAtlantic

Preliminary Results ROPOS DiscoveryCruise 2007

4 ConservationofDeep-SeaCoralsinthe Newfoundland andLabrador Region,

4.2.1Scope ofResearch

4.3.1 Why Are CoralsImportant?

ConservationProgress in EasternCanada 4.4.1 MaritimesRegion 4.4.2NewfoundlandandLabrador Region

NAFO RegulatoryAreas on theHighSeas 4.5 Toolsand StrategiesTo ProtectCorals and Sponges

PriorityAreas for Conservation 4.6.1Hatton Basin(National Waters) 4.6.2LabradorShelfEdgeandUpperSlope (National Waters) 4.6.3 Orphan Spurand Tobin'sPoint(National Waters) 4.6.4Flemish Pass(InternationalWaters)

4.7 Conservation ChallengesForNewfoundland andLabrador 4.7.1 PublicAwareness and Understanding....

4.7.2TraditionalManagement Styles and Advancements 4.8 Conclusions and ClosingThoughls....

APPENDIX 1:Systematic List of thePhylumCnidaria:Class AnthozoaNewfoundland

LiteratureCited Appendix 1

APPENDIX3:Dislribution MapsandDala used in the ProductionofMapsfromChapter 2 (Wareham&Edinger, 2007) and Chapter 3 (Wareham, 2009) APPENDIX4:ColieclionProlocolsforcoralsandspongesforNewfoundland,Labrador,

LIST OF TABLES

Table 1.1.Summary of growth rates and associated studiesforsomedeep-seacoral species.RG=RadiaIGrowth;AC=AxiaIGrowth;EA=EstimatedAge;TL=Total Length.·:SamplesfromNorthwestAtlantic

Table 2.1.Summary of coral occurrences by species from DFO Multispecies Stock AssessmentSurveys(RV)2003-2006,theNorthernShrimpStock Assessment Survey (RV)2005,records (FOPr)and samples (FOPs)from fisheries observations April 2004-January2006 (see Appendix 3for electronic version).Note:()=species suspected but not confirrned

Table2.2.Summary of coral frequencies by target fishery,geartype,andaverage depthsfished;datafrom fisheriesobservations documentedbetweenApril 2004 andJanuary2006.Note:GN=gilinet,LL=longline,CP =crab pot,OT

= Ottertrawl, ST=shrimptrawl,n=twin trawl,andTn= triple trawl...68

Table2.3.Summary of coral frequencies by species,target fishery,gearclass,andgear type;datafromfisheries observations documented between April 2004 and January2006.Note:GN=gilinet,LL= longline,CP=crab pot, OT=Otter trawl,ST=shrimp trawl,

n

Table3.1.Summary of coral frequencies used in distribution maps

Figure 1.1.Mapof studyarea highlightingbathymetricfeaturesolteasternCanada 5

Figure 1.2. Pholoof several gorgonian corals with calcifiedhoIdfasls:(left)Acanella arbuscula,and (right)Radicipesgracilis.Photoscourtesy ofDFOCanada.13

Figure1.3.Pholoof several solitaryscleraclinian cupcorals:(left)Flabellum macandrewii(Gray,1849) takenat 361minDesbarresCanyon,and(right) Flabellumalabastrum(Mosley1876)takenat946minHalibutChannel Photoscourtesy ofDFOCanada

Figure 1.4. Photosof coral by-catch fromtheHattonBasin area:(Ieft)Primnoa resedaeformisentangledinalrawl nelin2007,(right)Paragorgiaarborea from the NorthernShrimpSurvey in 2006. Photoscourtesy ofDFOCanada

Figure 1.5. Photos ofin situcoralhabilatsdocumenledon lheSouthwest GrandBanks (topleft)grenadier swimming wilhina Pennatulaseapenfieldin Desbarres Canyonal 900m;(lopright)close-upof Umbellulaencrinus(Linnaeus, 1758) colonyinDesbarresCanyonal 1657m.Notethesmallmysids hovering between the polyps;(bottom righl)Acanlhogorgiaarmatacolony withashrimp resting wilhin anAcanellaarbusculacolony;(bottomleft) redfishandspottedwolfish,lisledaslhrealened(COSEWIC,2001) resting around a small bouldercQveredwith corals,including: Keratoisisornata, Anthomastusspp.,Acanlhogorgiaarmala(Verrill,1878),neptheids,and sponges.Note injtialobservationshowedtwo redfishon either side ofthe

boulderwiththespottedwolffishcurledaroundthebaseoftheK.ornala colony.Photoscourtesy ofDFOCanada

Figure 1.6.Illustrationof an otter trawl used intheNewfoundland andLabradorregion

Figure 1.7.IIlustrationsoftrousertrawl,twintrawl,andcomponents used inthe NewfoundlandandLabrador region(DFO,1997) Figure 1.8.Illustrationsof separator grates and components(DF0,1997)

Figure 1.9.Illustrationof crabpotgear configurationused inthe Newfoundland and Labrador region(DFO,1997)

Figure 1.10.Illustrationsof gillnetconfigurations and components(DFO,1997) Figure 1.11.Illustrationof longline gear configurationandcomponents(DFO,1997) 26

Figure 1.12.PhotoofPrimnoaresedaeformis skeleton encrusted with colonial sea anemones,hydroidsand other organisms

Figure2.1.Deep-seacoralspecimenscoliectedoffNewfoundland,Labrador,andBaffin Island.OrderPennatulacea:A.)Dislichoptilumgracile;B.) Pennatula phosphora;C.)sea pen sp.6(?);D.)Pennlulasp.(?);E.)Pennatulagrandis;

F.)Umbellula Iindahli;G.)Haliplerisfinmarchica;H.)Anthoptilum grandiflorum;I.)Funiculinaquadrangularis.OrderAlcyonacea:J.)Radicipes gracilis;K.) Acanlhogorgiaarmala;M.)Acanellaarbuscula;N.)Paramuricea spp.(?);P.) Primnoaresedaeformis;Q.)Keraloisisornata;L.)Anlholhela grandiflora;0.)Paragorgiaarborea;W.)Anthomaslusgrandifiorus;X.) Gersemiarubiformis;Y.) Capnella florida.R.)OrderAntipatharia (?). Order

Scleractinia:S.)Desmophyllum dianthus;T.)Flabellumalabastrum;U.) Dasmosmilialymani;V.)Vaughanellamargaritata.Note:(?)=speciesnot

Figure2.2.Distributionofalcyonaceans inscientific surveys (RV)2003-2006and fisheriesobserverdata(FOP) 2004-2006

Figure2.3.Distributionofpennatulaceans,solitaryscleractinians,and antipathariansin scientificsurveys(RV)2003-2006andfisheriesobserverdata(FOP)2004-

Figure2.4.Coralspecies richnesspersetin scientificsurveydata.Mostspecioseareas were;A.)MakkovikBank-BellelsleBank,andB.)southwestGrandBank DistributionofcoralrichareasfromDFOsurveydataandfisheriesobserver data:C.)SoutheastBaffinShelf-CapeChidley,D.) Funk IslandSpur-Tobin's Point,andE.) FlemishCap

Figure2.5.Coral speciesrichness per set in scientificsurveydataandcoraloccurrences by Order fromfisheries observer data forMakkovik Bank-Belle lsieBank

Figure2.6. Coral speciesrichnessper setinscientificsurveydataandcoral occurrences by Order fromfisheriesobserverdataforsouthwest GrandBankandtailof

Figure2.7. Coral speciesrichnessper setinscientificsurveydataand coral occurrences byspeciesfrom fisheriesobserver data for SoutheastBaffinShelf- Cape

Figure2.8.Coral speciesrichness per set in scientific surveydataandcoraloccurrences by species from fisheries observer datafor Funk IslandSpur-Tobin'sPoint

Figure2.9.Coralspecies richnesspersetinscientific surveydataandcoraloccurrences by species from fisheries observer datafor FlemishCap

Figure3.1.Studyarea and sampling effort with distribution of deep-seaeorals highlighted.Datawas collected from;NorthernShrimp Survey (2005), New1oundlandandLabradorMultispeeiesSurveys(2000-2007),Aretie MultispeeiesSurveys(2006-2007),andfromfisheriesobserversaboard commercial fishing vessels (2004-2007)

Figure3.2.Distributionof deep-sea corals fromtheOrderAntipatharia(ineludes StauropathesarclicaLutken,1871,andBathypalhesspp.).Datawas collectedfrom;NorthernShrimpSurvey(2005),New1oundlandand Labrador MultispeeiesSurveys (2000-2007),Aretie MultispeeiesSurveys (2006-2007), andfromfisheries observers aboard commercialfishingvessels (2004-

Figure3.3.Distributionof deep-sea coralsfrom the OrderAleyonaeea(large gorgonians).Datawas collected from;NorthernShrimp Survey(2005), New10undlandand LabradorMultispeeies Surveys(2000-2007),Aretie MultispeeiesSurveys(2006-2007),andfromfisheriesobserversaboard commercialfishingvessels (2004-2007)

Figure3.4.Distributionofdeep-seacoralsfromtheOrderAleyonaeea(small gorgonians).Datawascollected from;NorthernShrimp Survey (2005), New10undland LabradorMultispeeies Surveys(2000-2007),Aretie

MultispeciesSurveys(2006-2007),andfromfisheriesobserversaboard commercialfishingvessels (2004-2007)

Figure3.5.Distributionof deep-sea corals fromthe OrderAlcyonacea(softcorals).Data wascoliectedfrom;NorthernShrimpSurvey(2005),Newfoundlandand LabradorMultispeciesSurveys(2000-2007),ArcticMultispeciesSurveys (2006-2007),andfromfisheriesobserversaboardcommercialfishing vessels (2004-2007)

Figure3.6.Distributionof deep-sea corals fromthe Order Scleractinia(solitarystony corals).Datawas collectedfrom;NorthernShrimp Survey (2005), NewfoundlandandLabradorMultispeciesSurveys(2000-2007),Arctic MultispeciesSurveys(2006-2007),andfromfisheriesobserversaboard commercialfishingvessels (2004-2007)

Figure3.7.Distributionofdeep-seacoralsfrom the Order Pennatulacea(seapens) Datawas collectedfrom;NorthernShrimpSurvey(2005),Newfoundlandand Labrador MultispeciesSurveys (2000-2007),ArcticMultispeciesSurveys (2006-2007),andfrom fisheriesobservers aboard commercialfishing

Figure3.8.Preliminary resultsofunique deep-seacoralhabitatsfromNewfoundlandand Labrador ROPOS Discovery Cruise(2007)includingclustersofKeratoisis ornatacolonies,termed'thickets',and large concentrations0fsea pens and

Figure3.9.Deep-seacoral species documentedin NewfoundlandandLabradorduring the2007ROPOS DiscoveryCruise;A.)Umbel/ulaencrinus;S.) Protoptilum

carpenteri; C.)Chrysogorgiaagassizii;D.)Flabellummacandrewi,and E.)

Figure 4.1.Mapof study areahighlightingimportantbathymetricfeatures offthecoast of

Figure 4.2.Map ofTheGullyMarine Protected Area(DFO, 2004a).Mapcourtesy of

Figure 4.3.Mapofthe NortheastChannel Coral ConservationArea(DFO,2002).Map courtesy ofDFO Canada

Figure 4.4.MapofLopheliaCoraIConservationArea(DFO,2004b).Mapcourtesyof

Figure 4.5.Mapof CAD-NAFO CoralProtection Zone(NAFO, 2007) with coral species richness.Richnesswasdetermined by the numberof coral species documented persetfrom DFO MultispeciesSurveys andfisheries0bservers

Figure 4.6.MapofNarwhat-CoraIProtectionZone(DFO,2007a)withcoralspecies richness.Richnesswasdetermined by the numberof coral species documented persetfrom DFO MultispeciesSurveys andfisheries0bservers from 2002 to 2007.Note the largecatch ofKeratoisissp. documentedin October 1999withintheexisting closure

Figure 4.7.Map i1lustratin9the Voluntary CoralProtection Zonewithcoralspecies richness.Richnesswasdetermined by the numberof coral speciesobserved persetfrom DFOMultispecies Surveys andfisheriesobserversfrom2002to

Figure.4.8.Map illustratingcoral and sponge by-catchesusingthresholds recommendedby theNAFOWGEBMfor internationalwaters and appliedto Research Surveydata (2005-2008) withinnationalwaters.Thresholds for largegorgonian by-catchare>2kgsand spongeby-catch threshoIdsare>

75kgs.Note:actualsetweightsforcoralsareiliustrated,however,actual sponge weights arenot,instead each point representsan occurranceof spongeby-catch>75kgs

Figure4.9.Map ofseamounts closedto bottom fishing within the NAFO RegulatoryArea

Figure 4.10.Location of candidate VMEs (ingreen) identified andrecommendedto NAFO by theWorkingGroup On EcosystemApproachTo Fisheries Management(NAFO,2008e),andactualareasdesignatedasinterimcoral protectionzones(inred)byNAFO,basedoncandidateVMEs(NAFO,

Figure4.11.Priorityareasforfuturedeep-seacoralresearchare highlighted ingreen, andprotectedareas arehighlighted inred

Figure.4.12. MapofHattonBasin,priority area for future deep-sea coralresearch. Map illustratesthecurrent Voluntary CoralProtection Zonewith coraIspecies richness. Richnesswasdeterminedbythe numberof coral speciesobserved persetfrom DFO MultispeciesSurveysand fisheriesobserversfrom2002to

Figure.4.13.Coralby-catchesfromHattonBasin:(left)Paragorgiaarboreaby-catchon trawl deck from Northern Shrimp Research Survey,and(right)Primnoa

resedaeformis fragments entangled ina commercial gill net.Photocourtesy

Figure4.14.MapoftheLabradorShelf,priorityareaforfuture deep-sea coral research Map illustrates coral species richness which was determined by the number of coral species observed per set fromDFOMultispecies Surveys and

Figure4.15.AlargeintactParagorgiaarborea colony collected justnorthofTobin's Pointduringthe2006DFOfalisurvey.Samplesofthisspeciesandsize,are relatively infrequent..

Figure4.16.MapofOrphanSpur-Tobin'sPoint,priority area for future deep-seacoral research.Map illustrates coral speciesrichnesswhich wasdetermined by thenumberofcoralspeciesobservedpersetfromDFOMultispecies Surveysand fisheriesobserversfrom2002to 2007....

Figure 4.17. Map of Flemish Pass,priority areafor future deep-seacorairesearch.Map illustrates coralspeciesrichnesswhich wasdetermined by the numberof coral species observed per setfrom DFO MultispeciesSurveysandfisheries

Figure4.18.Map of southwestGrand Banks,priority area forfuturedeep-seacoral research.MapiliustratesthecurrentCAD-NAFOCoralProtectionZone overlaid with coral species richness.Richness was determined by the number of coral species observed per set fromDFOMultispecies Surveys

Figure4.1g.Examplesofcoralsamplescaughtasby-catchbyalocal Newfoundland andLabradorgilinetfisher;(L-R)Primnoaresedaeformis, Paraborgia arborea,and Desmophyllumdianthusfrom MakkovikBank,LabradorShelf

Figure4.20.Examplesofcoralsamplescaughtasby-catchbylocalNewfoundlandand Labrador fishers;(L-R)Paramuriceasp.acquired off southern

Is OF ACRONYMS

Bedford Instituteof Oceanography

Centre ofExpertisefor AquaticHabitatResearch

CanadianBroadcastingCorporation CanadianCoast GuardShip Centre ofExpertise

Committee On Status Of EndangeredWildlifeInCanada DepartmentofFisheriesandOceansCanada

EcologicallyandBiologicallySignificant Areas

EnvironmentalNon-governmentOrganization EcosystemResearchInitiative

Fisheriesand Aquaculture Management

Groundfish EnterpriseAllocation Council General BathymetricChart ofthe Oceans

InternationalCouncilfor the Explorationof the Sea InternationalGovernanceProgram InternationalGovernance Strategy Integrated Taxonomic InformationSystem Local EcologicalKnowledge LargeOcean Management Area Memorandum ofUnderstanding

NorthwestAtlanticFisheries Organization

NAFO PotentialVulnerable MarineEcosystems Impactsof Deep-sea

NewfoundlandandLabrador'sExpanded ResearchonEcosystem- relevantbut Under-surveyed Splicers

Non-government Organization

NorthwestAtlanticFisheriesOrganizationRegulatoryArea

National ScienceEngineering ResearchCouncil

Regional FisheriesManagementOrganization

RemotelyOperatedPlatformfor Ocean Science

RemotelyOperated Vehicle

Significant AdverseImpact

UnitedNations General Assembly UNESCO-IOC Registerof Marine Organisms Vulnerable MarineEcosystem Vessel Monitoring System

Working Group onEcosystemApproachto FisheriesManagement WorldRegisterof Marine Species

Appendix1:SystematicListofPhylumCnidaria: Class Anthozoa Newfoundland,

Appendix 2:IdentificationGuideto Deep-SeaCoralsNewfoundland,Labrador,and

Appendix 3:Dataused in theProductionofDistributionMaps fromChapter2(Wareham

&Edinger,2007) and Chapter 3 (Wareham,200ga) on CompactDisk

Appendix 4: CollectionProtocolsfor Corals and Spongesfor Newfoundland,Labrador,

1 INTRODUCTION TO DEEP-SEA CORAL IN THE NEWFOUNDLAND AND LABRADOR REGION,NORTHWEST ATLANTIC OCEAN

Deep-seacorals(Phylum:Cnidaria),alsoknownascold-watercorals,arevibrantly- coloured animalsthat looklike planlsand comein variousshapes and sizes.First discoveredby deep-sea expeditions in thelate 17005(Pontoppidan,1755)theyare dredged fromthe seafloorfrom all overtheworld atalldepths(Freiwaldetal.,2004) Thestudy ofdeep-seacorals is costly and time-consuming duetothe great depthsof theenvironementtheyinhabitandtheassociatedlogisticsofconducting research in the

increasing due to the functionalrole(s) they play in providing habitat for otherspecies andcontributing towards habitatcomplexity(Robertsetal.,2009).Until recently,little was knownaboutthe distributionofdeep-sea corals in the northwestAtlantic with the exceptionof sporadicoccurrences from pioneering expeditions(Moseley,1881;Verrill, 1885;Agassiz,1888;Jourdan,1895;Pax,1932;Kramp,1942;Litvin & Rvachev, 1963;

Nesis,1963a;Nesis,1963b)withfewexploringNewfoundlandandLabrador waters (Jourdan,1895;Litvin&Rvachev,1963;Nesis,1963a;Nesis,1963b).The purposeof this thesis is to helpfillinformationgapson corals fromthe northwestAtlanticbyfulfilling three primarygoals

I Toidentifydeep-seacoralspecies andfrequenciesof occurrence;

" TomapthedistributionsofcoralsoffthecoastsofNewfoundland,Labrador,and

iii Todiscusspotentialconservationstrategiesandrecommendalternativeactions for national and regional resource managers

Chapter 1 introduces the thesis and includes;what are deep-sea corals,where they are found,whytheyareimpcrtant,andthreatstotheirexistence.lnaddition,background information on historical occurrences will be described.Subsequent chapters focus on basic biology of deep-sea corals,threats to deep-sea corals,andpreviouscoral

Chapter2 addresses two goals:

i. Toidentify deep-sea corals;and

This chapter was presented at the 3'dInternationalDeep-Sea CoralsSymposiumin Miami,Floridain 2005 and is published in a specialedition of theBulletin of Marine Science (Wareham&Edinger,2007)

Chapter 3 provides anupdate on the ongoing processofidentifyingandmappingdeep- sea corals in the northwest Atlantic,building onthe existing datasetdevelopedby Wareham andEdinger(2007).Collectively,it will be usedto identifyandhighlightareas forprotection within theNewfoundland,Labrador,and eastern Canadian Arctic regions Chapter3was published as Wareham (2009a) as part of Department ofFisheries and Oceans Canada (DFO)TechnicalReport summarizing researchconducted in the Newfoundland andLabradorRegion titled"The Ecology of Deep-sea Corals of Newfoundland and Labrador Waters:Biogeography,Life History,Biogeochemistry,and Ro/eofCrilicalHabitaf (Gilkinson&Edinger,2009)

Finally,Chapter 4 highlights progress on coral conservation todate,discusses management strategies to identify priority areas for future protection,and outlines

challengesthe NewfoundlandandLabrador Region faces.Thischapter concludes with recommendations foraddressingthesechallengesinordertosuccessfullyprotectcorals

FourAppendicesfollowthe thesis.Appendix1provides aSyslematic Lisl of Phylum

BaffinIsland,Canadaemphasisedin bold.Appendix 2isa"user-friendly"ldentificalion GuidetoDeep-SeaCoralsNewfoundland,Labrador,andBaffinlsland,Canada,in poster format(Wareham,200gb).I developed the poster,anditwas reproduced by

theposterisincludedonacompact diskinAppendix 3.Thecompactdiskalsoincludes Adobe PDFsofmapsand data usedin mappingof deep-sea corals pubIishedinChapter 2(Wareham &Edinger,2007) and Chapter3(Wareham,2009a).Appendix4,Collection Prolocolsfor Corals and SpongesforNewfoundland,Labrador,andBaffinlsland, Canada,containstwoexamples of standarddeep-sea coralcollectionprotocolscreated for this thesisandutilizedbyDFOstaff,and regional fisheriesobservers (SeaWatch

Thereare severallimitations regardingdatainterpretationpertaining toChapters2 and 3.Data used inthisthesiswere collected byDFOandby the FisheriesObserver Program(FOP).Bothsources sample from a variety of substratesusingdifferentgear types. Forexample,DFOresearch vessels sample with a CampelenTrawl usedon relatively levelsea floors and arelimited to depths<1,500m. EachDFOsurveysetis standardizedbydepthandtowduration(seeMcCallum&Walsh,1996). The needfor consistenttowspeed and distanceleadstoa bias that favours Ievel sea floor

environments, excluding steeper slopes and canyons. On the other hand,data from the FisheriesObserverProgramare derived from commercial vessels using a variety of gear types,representingpreferredfishingareasbasedonpastcatchrates and past experience of individual skippers.Observer dataincorporatesmanyfisheriesfrom a variety of depths, gear classes (i.e.mobile and fixed),geartypes(e.g.shrimptrawl,twin trawl),marinehabitats(e.g.steepcanyons),andseafloorsubstrates(e.g.boulders fields,mud,orsand).lnshort,DFOresearchdataarebiasedtowards'trawlable' substrates and observer data are biased towards preferredfishinggrounds

Thegeographic scope of this thesis incorporates a large portion0fthenorthwestAtlantic offthecoastsofNewfoundland,Labrador,andsoutheastBaffinlsland(Fig.1.1).lt encompasses theNorthwestAtlanticFisheriesOrganization (NAF0) fisheriesregulatory areadivisions;3KLMNOP(Newfoundland),2GHJK(Labrador),andOAB(easternBaffin Island). Datafor this thesis was gathered fromDFO researchsurveys(2003-2008),as well asfroma partnership with industry (Northern Shrimp Survey). Athird importantdata SQurce wasfromfisheriesobservers on board commercial fishing vessels operating withinthe NAFO regulatoryareas within Canadian jurisdiction All coraltaxa recordedduringthestudy wereincorporated intothe database.Basedon the Integrated Taxonomic InformationSystem (ITIS),taxawere documented fromthree subclasses: Octocorallia (;Alcyonaria),Hexacorallia(;Zoantharia),and Cerianthipatharia(seeAppendix1).Octocoralsinciudedtwoorders;pennatulaceans (seapens)and alcyonaceans (soft corals and gorgonians).Hexacorals included scleractinians(stonycupcorals);andCerianthipathariainciudedantipatharians(black- wire corals).Therewere no hydracoralsrecorded

ISLAND

CANADA

17 ATLANTIC OCEAN

Figure 1.1.Map of study areahighlightingbathymetric features off eastern Canada

Therearevariousother papers on deep-seacoralsto whichIhave contributed,butare not includedinthisthesis. Wehavedescribed methods forworkin9 with fisheries observers(Warehametal.,2007),andcoraldistributionsintheNAFOregion(Fulleret al.,2008).Wehaveanalysisedthedensitiesofcoralsandspongeswith associated fishing efforts in proximity toHanon BasininNAFOdivisions2G-OB(Kenchingtonetal., 2010;Wareham et aI., 2010).We have documented the ecological importanceof corals in Newfoundland andLabradorwaters to fisheriesspecies (Edingeretal.,2007b),and fisheriesimpactsuponcorals(Edingeretal.,2007a).WehavedescribedAdenosine triphosphate and lipid biochemistry of severalspecies ofcorals(Hamouteneet aI., 2008a,b),andtissue stable isotope geochemistry of11species of deep-sea corals from AtianticCanadianwaters(Sherwoodetal.,2008)

Corals are simple animals,referred toindividually aspolyps(Birkeland,lg96;Ruppertet al.,2004;Hopley,inpress).Eachpolypiscomposedofaring(s)0ftentaclesusedfor capturingfood,amouth foreating, and atube(actinopharynx orstomadem)leadingtoa central gastrovascular cavity (coelenteron)for digesting food. Corals canbesolitary,

known tropicalreef·buildingvarietiesarefoundin shallower warmerequatorialwaters Theyare restrictedtothe photic zone becausethey haveendo-symbiotic relationships with algae and arereferredto as zooxanthellate corals.Thecoralcolony(:corallum) providesthesubstrateforalgae to live inwhiletheyphotosynthesize energy from the sunand convertitto food.Inexchange,the algae excrete metabolicwaste whichis recycledas food bycoralpolyps(Muscatine&Porter,1977;Hallock& Muller-Karger,

Incontrast,deep-seacorals arenotas well-knownbecausetheyarerarelyseen (Robertsetal.,2009).Mostspeciesarefoundbelowthephoticzone.Deep-seacorals arereferredtoasazooxanthellatecorals,whicharestrictlysuspensionfeeders.They depend entirelyon currents and otheroceano9raphicprocessestotransport food to them,likezooplankton and detritus,which has beenimported from thewater column nearthewatersurface(Freiwald,2002;Sherwoodetal.,2008).Currentsnotonlycarry foodbut alsoprevent accumulation of silt,which can smotherthepolyp(Robertsetal., 2009)

Basedonanatomydeep-seacoralscanbedividedintoseveralmaingroups:octocorals, scleractinians,andantipatharians.Stylasteridae andZoanthidaearetwo additional 9roups(Cairns,1992;Ocana&Brito,2004) but werenot foundwithinthestudy area, therefore are notcovered.Octocorals(ClassAnthozoa:SubclassOctocorallia)include the90r90nians,soft corals,andpennatulids(seapens).Thisgroupiseasily distinguishable.Eachpolypconsists of eighttentaclesallcontainingsc/erites-special internal structures constructed of calcitic calcium carbonate (Robertsetal.,2009) Almost allspeciesin thisgroup are colonial withtheexception of onegorgonian(Bayer

&Muzik,1976). Gorgonians haveahardor consolidatedinternalskeleton whichis constructed of either proteinaceous gorgonin, calcium carbonate(calcite or aragonite) or amixtureof the two (Bayer,1973).Softcoralshaveahydroskeletonand relyon water andhydrostaticpressuretomaintaintheirshape(Fabricius&AIderslade,2001) Similarly,sea pensmaintain theirshapeusing hydrostatic pressure butbenefit froma centralinternalcalcareous axialrachis,foradded support andmovement(Williams,

Scleractinians(ClassAnthozoa:SubclassHexacorallia),alsoknown asthestony corals, canbe foundas colonial~reef-builders"or solitary cup corals.Mostdeep·sea species

arethe latterwith few exceptions (Cairns, 2007). Stonycoralsare easilyidentifiedby theireXDskeletonsconstructed of aragonitic calcium carbonate

ThefinalgrouparetheAntipatharians(ClassAnthozoa:SubclassCeriantipatharia),also known astheblack-wire corals.Members ofthisgroup are all colonial.Polypsare simple,containing six tentacles andlackingsclerites.Theyhave auniqueinternal chitinous skeletonforstrength and support,whilemaintaining flexibility.Theskeleton is cDveredwithtinyspines.Taxonomicidentificationofthisgroup is difficult. Species identification requiresclose inspection of polyps (size and structure),spinesand skeletonmorphology(Goldsbergetal.,1994;Opresko,2002;Molodtsova&Budaeva, 2007)

Coralsmay be the longest livinganimals onEarthwith some specie5reachingseveral thousandyearsinage(Roarketal.,2006;Robertsetal.,2009).Growthratesformany deep-seacoralsareunknown,howeversomespecieshavebeeninvestigated(Table 1.1);PrimnoaresedaeformisGunnerus, 1763(Risketal.,2002;Sherwoodetal.,2006), KeratoisisornataVerrill,1878(Roarketal.,2005;Sherwood&Edinger,2009), DesmophytlumdianthusEhrenberg,1834(Risk et aI.,2002),Hatipteriswillemoesi (Kblliker,1870;Wilsonetal.,2002),Chrysogorgiaagassizii(Verrill,1883;Vinogradov, 2000)andStauropathesarcticaLOtken,1871(Sherwood& Edinger,2009).All have extremely slow growth rates and some species can surpass 100'50fyearsinage (Sherwoodetal.,2006;Sherwood&Edinger,2009)

Table1.1. Summary ofgrowthrates and associated studiesforsome deep-sea coral species.RG=RadiaIGrowth;AC=Axial Growth;EA=Estimated Age;TL=Total Length

CoralSpecies GrowthRates & Estimated Longevity References Primnoaresedaeformis 'RG = 83 ±6t0215 ±37~myr' Sherwood&Edinger,2009;

'AG=1.00±0.09t02.61±0.09cmyr' Sherwoodetal.,2006 (gorgonian) 'EA=18tol00yr(2009)

'EA=<700yr(2006!

Sherwood&Edinger,2009 Acanellaarbuscula 'RG - >20~myr'

(gorgonian) 'AG=>0.30cmyr' 'EA =30yr(growth nng counts!

Keraloisis,Isidella,or RGR- 50·160~myr' Acanellasp.(gorgonians) (Alaska,USA)

Sherwood&Edinger,2009 'RG - 53±9to 75 ± 11~myr'

'AG=0.93±0.08cmyr' Keratoisisomata 'EA=94±7t0200±30yr

RG - 50-110~myr' (gorgonian) (Davidson Seamount,offCalifornia,USA!

RG -111~myr'(Tasmania! Thresheretal.,2007 RG = 130· 290~myr'(NewZealand) Traceyetal.,2007

Paragorgiaarborea AG-0.8-4cmyr'

~~~:~=nai ~~~~;

(gorgonian) (NewZealand&Norway)

Mortensen,2005 '1.62 ± 0.22 cm vr' Sherwood&Edinger,2009 Paramuriceaspp 'RG-92 ± 18~myr'to Sherwood&Edinger,2009

205 ± 20~myr' (gorgonian) 'AG = 0.56 ±0.05~myr'to Chrysogorgiaagassizii 0.58±0.08cmvr'

Vinogradov,2000 AG-lcmyr'

!gorgonian) ~~:lIbasedT~:°lii~;~ ~~wth!

colonies AG=3.9±0.2cmyr' Halipteriswillemoesi MediUnl

~t~~;

~~!~

(sea pen) colonies AG=6.1±0.3cmyr'

~~~;~es ^~t:

^~~75c~S

Sherwood&Edinger,2009 Siauropalhesarclica 'RG - 33 ± 11t066± 11~myr'

(antipathanan) 'AG=I.22±1.46tol.36±0.20cmyr' 'EA=55±8t082±31vr Leiopalhesglaberrima RG-0.0145mmyr'

(antipatharian) EA= 200yr(Flonda,USA) RG~10~mvr'

Investigationsinto thehistologyof soft corals indicated that they too may exhibit slow growth rates especially in early stages of recruitment (Cordes et al.,2001;Sunetal., 2010).Forexample,newly settled Drifa sp.reached only 5 mm linear length in 7 months, while Duva florida exhibited no branching of polyps in 11 months,and Gersemia fruticosareachedonly 10 mm linear length in 7 months (Sun et aI.,2010)

1.1.5 Reproduction

Most knowledge of coral reproduction is from tropical species with very little known aboutdeep-seaspecies.Sexualreproductioncanbeeitherhermaphroditicwithmale (spermatocysts) and female (oocytes) gametes located on the same colony,or gonochoristic with male and female gametes located ondifferentcolonies.Gonochoristic species,alsoknownasunisexual,produceplanulalarvaewhichcan developfrom internal or external fertilization.Internalfertilization resultsineggsbeingfertilizedand developed withinthe maternalcolony, known as brooding (Richmond&Hunter,1990) Externalfertilizationresultsin eggs being fertilizedanddeveloped withinthewater column,knownas broadcast spawning (Richmond &Hunter, 1990)

Hermaphroditicspecies produce planulalarvaeaswell,butspermatocysts and oocytes candevelopondifferentlocationswithinthesamecolony(i.e.mesentery,poIYP,or colony),ormaydevelopatdifferenttimeperiodswithinthesamecolony (Rinkevich&

Loya,1979).Thisdevelopment of gametes can occur simultaneously orsequentiallywith oocytesdevelopingfirstfollowedbythedevelopmentofspermatocysts(Rinkevich&

Loya,1979).Oncefullydeveloped,planulalarvaearereleasedintothe water column from the parent as mature planulae (Richmond &Hunter,1990;Fabricius& Alderslade, 2001)

In Newfoundlandwaters reproductive biology offourspecies of soft corals (neptheids) hasbeeninvestigated(Sunetal.,2010).Drifasp.wasfoundtobea hermaphroditic internalbrooder.Drifa glomertawas found to beaninternalbrooderbutitis undetermined whether it is gonochoristicorhermaphraditic.Methodof reproduction of twootherspecies,D.f1oridaand G.frulicosa,wasnotdetermined(Sunetal.,2010)

Manygapsinourunderstandingofthegeneralbiologyofdeep-sea coralsremain.There islittleinformationavailableonageofmaturity,fecundity,reproductionandrecruitment, resilienceandresistance todamage, andratesofrecovery(Robertsetal.,2009)

1.1.6Where Do Deep-SeaCorals Live?

Deep-seacorals areusuallyfoundinareas with pronounced bathymetric reliefsuch as deep-seacanyons, seamounts, and along the continentaledge,slope andrise (Deichmann,1936;Nesis, 1963b;Tendal,1992;Breezeetal.,1997;Macisaacetal., 2001;Mortensen &Buhl-Mortensen,2004;Gass & Willison,2005;Bryan & Metaxas, 2006;Wareham &Edinger,2007;Wareham, 2009a).InNewfoundlandandLabrador, deep-sea corals can be found on the continental shelfandedge<2OOm,andonthe continental slope between 200-2,000m deep(Nesis,1963ab;Gass&Wiliison,2005;

Mortensenetal.,2005;Wareham &Edinger,2007;Wareham,2009a),with some speciesdocumentedonthecontinentalriseatdepthsof2,200m(Bakeretal.,2008;

Substratepreferencesare species-specificbasedon availability of hard substrates as wellasthe physiologyofindividualspecies. Substrates can varyfromabiotic (e.g boulders, cobbles,pebbles,and mud),tobiotic(e.g.othercorals,bryzoans,sponges, andlivinggastropods),toevenanthropogenicsuchasabandonedfishinggear,and plastics (Wareham &Edinger,2007;Bakeretal.,2008)

ROPOSDiveR1072).Othercoralssuchassolitaryscleractiniancup corals (e.g Flabellumspp.)simplyreciineontheseafloor(Fig.1.3)withnoattachmentor anchored appendage (Mortensen etal..2006; Wareham&Edinger, 2007; Baker etal..2008)

Figure1.2.Photo of several gorgoniancoralswithcalcifiedholdfasts:(left)Acanella arbuscula,and(right)Radicipesgracifis.Photoscourtesy ofDFO Canada

Figure 1.3.Photo ofseveralsolitaryscleractinian cupcorals:(left)Flabellum macandrewii(Gray,1849) taken at 361 minDesbarres Canyon,and (right)Flabellum alabastrum(Mosley1876)takenat946m in Halibut Channel.PhotoscourtesyofDFO

Mostcorals inNewfoundland and Labradorwaters werefound at depthsgreaterthan 200m(seeChapter2,Table2.1).lthasbeenpostulatedthatthispossible restriction to deeper water may be due to several environmentalfactors such asstrongcurrentsalong the edgeofthecontinental slope,suitable substrates,andconstant temperature ranges

Mostcoralsdependentonoceancurrentsandotheroceanographic processes (e.g upwelling,and gyres)todeliver particulate organicmattersuspended in the water column (Moore &Bullis,1960;Tendal,1992;Bryan& Metaxas,2007),and winnow away fine sediments (Wainwright &Dillon,1969).Most speciesneedsuitablehardsubstrates for attachment (Mortensen &Buhl-Mortensen,2005).OntheScotian Shelf,deep-sea corals prefertemperaturesthat range between 3.5-13°C withhightemperatures most likely limitingdistribution (Mortensen etal.,2006),although ithas been found that some speciesofsoftcoralscantoleratetemperaturesaslowas-1°C(Cimbergetal.,1981;

Freiwald,2002).However,the general distribution (i.e.onbanksvs.edge and slope) of deep-sea coralsintheNew10undlandandLabradorregion may beIimitedlargelybycold watertemperatures.Thecoldintermediate layerisafunction 0ftheLabradorCurrent and brings sub-zero waters to theNew10undlandand LabradorShelvesdownto-200 m(Dunbar,1965;Petrieetal.,1992).Asaresult,thismay restrict distributionstodeeper waters onthecontinental slope where temperatures are more stable and often warmer, comparedto bank tops where temperatures canfluctuate(Nesis1963b).Some gorgoniansintheNewfoundlandandLabradorregion,likePrimnoaresedaeformishave beenfound as shallow as 162monSaglekBankoffnorthernLabrador(Wareham&

Edinger,2007).Thisarea in particular isknown forstrong currentswhich drain from Arctic waters viathe HudsonStrait andDavisStrait (Piper,2005). Suchlarge influxesof cold waternotonly provide suitable substrates butdeliverfoodandoxygenaswell.More importantly,theymaintaina constanttemperatureof cold water at much shallower depths than other areas withinthe region

Bryanand Metaxas (2006) observedthatPrimnoa resedaeformisand Paragorgia

where optimal water temperaturesranged5.1- 9.0°e,along with other environmental

factors.Otherstudiesfromthe Scotian Shelf suggestthat high temperaturesarea limiting factorfordeep-seacoral distributions (Mortensen et al.,2006).However,both studiesmay not fit the NewfoundlandandLabrador regionwhere10wtemperaturesare morelikely tobealimitingfactorforcoraldistributions,particularlyon bank tops«200 m;Edingeretal.,2007b)

There isanon-going debate overthe relative importanceofsubstrateversus oceanographicinfluences(e.g.currentstrength,temperature)ingoverningthe distributions of deep-sea corals (Nesis,1963b;Bryan&Metaxas,2006;Mortensenetal., 2006).Cinbergetal. (1981)usedannualmean temperature andsubstrate as predictors of coraldistributions.Nesis(1963b)linkeda change in seatemperaturewith changes in species assemblages,while Mortensen et al. (2006)found thatsubstrate and temperaturearethe mostimportant variablesinfluencingcorals distributions.Towhat degree specific environmentalfaclorssuch astemperature,slope,substrate,salinity, andchlorophyli ainfluencethe presence or absence of corals is nottrulyunderstood Bryanand Metaxas(2007)used known corallocationsand severalenvironmental parameterstodevelopa predictivemodel forcoral distributianS.How8ver,the inappropriatebathymetric scaleused intheirmodel precludeddefinitiveconclusions (Etnoyer&Morgan,2007). Asmoreenvironmentalinformationbecomes available and betterpredictivemodelsaredeveloped,keyfactorsinfluencingthedistribution of corals can beincorporated into predictive modelsandtested interms0ftheirrelative

1.1.7Ecologicallmportance

Deep-seacoralsnotonlyliveinbenthicecosystems,theyareimportantfunctional components oftheseecosystems.Their presence providestructure,adds structural and

biological complexity to the deep-sea,and create micro-habitats for other species (Austeretal.,2005;Buhl-Mortensen&Mortensen,2005;Auster,2007;Etnoyer&

Warrenchuk,2007;Mooreetal.,2008;Bakeretal.,2008).SpecieslikeParagorgia arboreaare consideredtobe one ofthemost important habitat-formingdeep-seacoral speciesbecauseof its'large size,reachingupt03m inheightoff eastern Canada (Mortensen &Buhl-Mortensen,2005) and up to10m offNewZealand (Smith, 2001) Denseconcentratioins of corals have been referred 10 as'coral-gardens'off the Aleutian Islands(Cimbergetal.,1981;Krieger&Wing,2002;Stone,2006),'gor90nianforests'on theScotian Shelf (Lees,2002;Mortensen &Buhl-Mortensen,2005),and'sea penfields' offNewfoundland andLabrador(Wareham,2009a).'Gardens', 'forests',and'fields'refer to large concentrations of corals with high species diversity0rbiomass abundance (Breezeetal.,1997;Watiing&Norse,1998;Kreiger,2001;Lees,2002;Freiwaldetal., 2004;Mortensen &Buhl-Mortensen,2005).Highby-catchratesandfisheriesobserver photosindicate'gorgonianforests'may exist off Cape Chidley,Labradorin close proximityto Hatton Basin(Macisaac et aI.,2001; Gass & Willison,2005;Wareham &

Edinger,2007;Wareham,2009a;Warehametal.,2010;Fig.1.4)

combined withrelativelysmall dataset (2 years).Otherstudieshaveshown stronger resultsduetodirect observationsusinggillnets(Huseb0 etal.,2002) and video observations(Austeretal.,2005;Costelloetal.,2005;Mortensenetal.,2005)

Figure 1.5.Photosof in situ coralhabitatsdocumented on the SouthwestGrandBanks (topleft)grenadier swimming within a Pennatu/asea penfieldin DesbarresCanyon at 900m;(topright)close-up ofUmbeliu/aencrinus(Linnaeus,1758) colonyin Desbarres Canyon at1657m. Note thesmall mysidshoveringbetweenthepolyp5;(bottomright) Acanthogorgiaarmata colony with a shrimprestingwithin anAcanellaarbusGulacolony;

(bottomleft)redfishandspottedwolfish,listedasthreatened(COSEWIC,2001)resting around a small bouldercQvered with corals,including:Keratoisisornata,Anthomastus spp.,Acanthogorgiaarmata(Verrill,1878),neptheids,andsponges.Noteinitial

curled around the base of theK.ornatacolony.Photoscourtesy ofDF

a

1.2.1BottomFishingPractices

Therearemanyanthropogenicthreatstocoralsbutthemostprevalentarebottom fishing practices andfishinggear that come in eontaetwiththe seafloor(Probert,1997;

Watling& Norse.1998;Fossaetal.,2001;Hall-Spenseretal.,2002;Grehanetal.,

followingseetion willdescribebottomfishinggear types used intheNewfoundlandand Labradorregionand discuss how eachimpactsdeep-sea corals.0therthreatsare diseussedaswell,buttoalesserdegree

Bottom fishinggear can be divided into mobile andfixed-gearclasseS.Mobilefisheries aetively pursue thetargetspecies and ean involvetrawling,dredging,or seining (Figs 1.6&1.7).Fixedgearfisheriesusea sit-and-waitstrategy,where the gear is positioned inonelocation withthepurpose of entanglement,entrapment,0r hookingthetarget speeies.Deepwaterfixedgearfisheries in NewfoundlandandLabradoruseerabpots, longlines,andgillnets(Figs.1.9-1.11)

Trawling,alsoreferredtoasdragging,isthe most widely-used fishingmethodinthe northwestAttantie(Fulieretal.,2008).Thereareseveraltypesoftrawlsused in the NewfoundlandandLabradorregion;otter (Fig.1.6),twin(Fig.1.7), triple,and shrimp Trawlinginvolvesthe dragging ofalargenet aeross the seafloor. The mouthof thenet is heldopen bythe forwardmotion ofthevesselcombined with the spreadingaetionof thetrawldoors as wellas floats positioned alongthe headline(Fig.1.7).Whilethebasic principle oftrawling has notchanged sinee it's ineeption inthe14^Iheentury(March,

1953),technolo9icaladvanceshaveimprovedcatchefficiencyandadaptabilityoftrawls tooperate in a variety ofhabitats

~

Groun<lr"" =.~ """' -

~

M.... IIIrI II

011110001 TrouserTrawl

~ ^~

Figure1.7.llIustrationsoftrousertrawl,twintrawl,andcomponents usedinthe NewfoundlandandLabradorregion(DFO,1997)

SEPARATOR GRATE

SYSTEMS ^panel

Figure 1.8.Illustrationsof separator grates and components (DFO,1997)

Trawling hasbeen comparedtoploughing afieldorclear-cuttingaforest(Watling&

Norse, 1998Anderson&Clark, 2003).Once trawled impactedareasbecomes more homogeneous throughmortalities and removal ofbioticcomponents such aslarge benthicmegafauna(e.g.corals,hydroids,sponges).TrawlingaIso alters abiotic

components suchasboulders,sand and mud-dominatedenvironments(Auster&

Langton,1999;Hall-Spenser,2002; Mortensenetal., 2005;Edingeretal.,2007a).Trawl doorscan weigh in excess of one ton each and createdeepfurrows in soft substrates as theyare draggedduringnormal use (Roberts,2002;Watling,2005).It hasbeen shown thattrawldoors can impact and destroy certain infaunal taxa (Gilkinsonetal.,1998) Experimentalstudies carried out in eastern Canada have shown thatthe combined effects of trawling can cause damage and mortality to epifauna and infauna (Prena etal., 1999:Kenchingtonetal.,2001;Gordon etal.,2003) and change sedimentstructural properties (Schwinghameretal.,1998)

Commercialtrawlingis carried outona large spatial scale with preferred fishing areas repeatedly fishedas seen onthe Grand BanksofNewfoundland(Kulka& Pitcher,2002;

DFO,2006).lndividualtowscanbeconductedover1-10hrs.Theoverallfootprintcan bevast,particularlyon an accumulativebasis.However,thedegreeofimpactwillvary dependingonthe biota present in the area beingtrawled.For example, instructurally complex coralhabitats, the initial pass is the mostdamaging(Kriger&Wing,2002;

Anderson & Clarke,2003;Rice,2006)resulting in largequantitiesofcoralby-catch (Probertetal.,1997;Kreiger,2001)

Othermobile geartypessuch as dredges canhavesimilar effects0ntheseaflooras trawling.Dredgestarget soft sediment habitats and can damage infaunal communities (Gilkinson et aI.,2003) as well asmegafaunacommunities (Veale etaI.,2000;Thrush&

Dayton,2002)

Fixedgear fisheries (e.g.crab pots,gillnets,longlines) have aIsobeen shown to capture anddamage deep-seaccrals(Huseb0etal.,2002;Mortensenetal.,2005;Wareham &

Edinger,2007).Althoughfixedgears are stationary, spatial coverage can still be significantbecausethese gears are often linked.IntheNewfoundlandandLabrador region,crab pots arelinkedtogether as a string of baited traps,called'strings'or'f1eets', withupt050pots perfleet(Fig.1.9).Impactsoccur whenthe fleetisretrieved,causing thecrab potstobe dragged across the seafloorwheretheycan ensnarland entangle

Figure 1.9.IIlustrationofcrabpotgearconfigurationusedinthe Newfoundlandand Labrador region(DFO,1997)

Bottomgillnets operateunder thesame principle,andean becomprisedof many panels (91.6mper panel) strung together withupto 70 panels perfishingset(Benjaminsetal., 2008).Howthe netis positionedinthewater column depends on thetargetedspecies Forsemi-pelagicfishes,theglllnethangs in thewater column like a giant wall nearthe seafloor(Fig.1.10).Thetop of each gillnetpanelis outfitted with floats (floatline)and the bottomwithlead-rope(leadline).In NewfoundlandandLabradorgroundfish fisheries,somefishersset bottom gillnets withno floats,which allowsthe panelsto bunch together vertically for purposes of entanglingthe targetspecies(W.DeGruchy, SeaWatchLtd.,personal communication,Sept.5,2007)-thismethodisnotillustratedin Figure 1.10.Regardlessofhowthe net is positioned,gillnetshavebeenshownto

capture and damage corals (Mortensen et aI.,2005;Gass&Willison,2005; Wareham&

Edinger,2007)

Figure1.10.llIustrationsofgilinetconfigurationsandcomponents(DFO,1997)

Longlineisanothertypeoffixedgearusedtotargeteitherpelagicorbenthic species Bottomlonglines are seton the seafloorwith amainlineconsistingofhundredsofbaited hooks,calledgangions.Eachendofthemainlineisanchoredtothe bottom,and marked atthesurfacewithbuoysand'hi-fiyers'(Fig.1.11).Asalonglineisretrievedthemainline becomes tautcreating a'clothes-line'effectacrossthebottom.Coralsin thepath ofthe longlinewillmostlikelybetilted,entangled,removed,ordamagedduring theretrieval process (Mortensen etal.,2005).This isparticularly significant forlargegorgonian coralsthatneedtomaintainanuprightposition.lfacolonyisdamaged (e.g. branches severed)itmay become more susceptible to parasitic organisms suchashydroids,or

colonialsea anemones (Fig.1.12),whichhas beenobservedinAtianlicCanada (Mortensenetal.,2005;Wareham&Edinger,2007)

Figure 1.11.Illustrationoflonglinegear configuration and components(DFO,1997)

Figure1.12.PhotoofPrimnoaresedaeformisskeleton encrusted wilh colonial sea anemones,hydroids and other organisms

Itwas surprising tofind how frequentlycorals were capturedbyfixedgearsinthe NewfoundlandandLabradorregion (Wareham&Edinger,2007).Thisismostlikelydue toa highercatchability of corals in certain areas that are targetedby fixed gearfishers Fixedgearfisheriesoperating on the sQuthwestGrandBankstargetareasthatare considered 'untrawlable'such as steep canyon walls and areas withrockysubstrates Theseareashave not beenimpactedto thesamedegreeas'trawlable'areas,andasa result,willmost likely havea greater abundance of corals and a greater chance of catching them

Whileboth mobileand fixedbottom gearscatch corals,theimpacts of trawling pose the

benthicpopulations, communitystruclure, andhabitatsbyremQvinglargemegafauna andalteringphysical componentsresultingin alossofbiodiversityandhabitat complexity on alargescale (DFO.2006)

Inadditionto bottom fishingpractices,thereare otherthreatsto deep-sea corals, including: hydrocarbonexploration,bio-prospecting,scientificresearchsurveys,ocean acidification, submarine cables,and aquacultureactivities

1.2.2.1HydrocarbonExploration

Hydrocarbonexplorationhasexpandedsince the 1980'sin the Newfoundlandand Labrador regionwiththreeproductionfieldsoperating onGrandBank;Hibernia(1979), Terra Nova(1984),andWhiteRose(1984) oil explorationplatforms(see oil fields inFig 1.1). Exploration continues toexpand with several projectsunderdevelopment

(Hebron/BenNevis,WhiteRose Extensions, HiberniaSouthExtensions and GardenHill South;NewfoundlandandLabrador,2010).Hydrocarbonexplorationisconsidered a threat to deep-seacoralsthrough the discardingoffine drill mud,aby-productofthe drillingprocess (Raimondi etal.,1997; Colmanetal.,2005).Thesemudscan accumulateneartheplatformorbetransportedfutheraway(e.g.upto1km)before settling(Neff,1987)andareconsidereddetrimentaltosuspension-feederslikecorals because itcan accumulate on polyps andinhibit feeding(Dodge etal.,1974;Dodge&

Vaisnys, 1977;Dodge&Lang, 1983).Whilemost research hasbeen carriedouton tropicalcorals,more recentlyLophelia pertusahasbeen observed growing on oil platforms in the NorthSea (Bell & Smith,1999; Roberts,2002; Gass&Roberts,2006), which wouldindicate thatsome speciestoleratesomedegreeof exposuretodrillmuds HOW8ver,the NorthSea platforms were notfixed tothe oceanfloor,therefore, colonies weremostlikelyisolated from drillcuttings which wouldhaveaccumulated onthesea f1oor.Ontheotherhand,platformslikeHiberniaarefixedtotheseafloor,thereforemay impactsoft corals which are found onGrand Bank in thevicinity of the oilfields (Wareham& Edinger,2007).The long-termeffect of discarded driIIcuttings on coralsis

1.2.2.2Bio-prospecting

Bio-prospecting is theharvesting of biological organismsforscientificandcommercial purposeswiththelatterinciuding;production ofpharmaceuticaIdrugs(e.g. anti-cancer), cosmetics(e.g.facecreams),nutritionalsupplements(S.Pomponi,FloridaAtiantic University,personal communication,Dec.5,2005),andmaterialsusedinbone reconstruction(Ehrlichetal.,2006).Bio-prospectingposesathreatto deep-sea corals becauseitphysically removes whole or parts of coralcoloniesfrom theseafloor:but more importantly there arenolegalmanagement regimes in place toregulate or policeit

in the deep-sea.Currently,there isnoofficial bio-prospectingoccurring withinthe Newfoundland andLabrador region,whichIam aware of

1.2.2.3ResearchSurveys

Aless-recognized threat todeep-sea corals isthe destructivenature ofbottom trawls, anddredges(Jennings &Kaiser,1998)currentlybeing used by DFO multispecies surveys.Duration ofscientific trawlsis relativelyshort (15minutes)comparedto commercialtrawlsets (1-10hrs)but continueto impact benthicenvironments.Modern soft-touch exploration technologies are available suchas ROVsandhave been usedin Canada (Haedrich&Gagnon, 1991;Mortensenetal.,2000;Wareham,2009a) Unfortunately,soft-touchresearchtoolsarecostty(i.e.operationalcosts,expertise),and fundsarenot readily-available

DepartmentofFisheriesandOceansScienceBranches inAtlantic Canadahavefunded andcontinuetofunduniqueresearchopportunitiesutilizingaspecializeddeep-sea ROV,calledRemotely Operated Patform for Ocean Science(ROPOS). Carrying out such scientific endeavours takesyearsto plan,but theoutcomesarebeneficial,with the abilitytocarryoutdiverseresearch inavariety ofhabitats inthedeep-sea(Bakeretal.,

1.2.2.4Climate Change

Climate changeincludes global warming and oceanacidification.Globalwarming is causedby greenhousegasesreleased into theatmosphere.Thesegasses create a blanket effect,which cause global surfacetemperatures to riseincludingmean-sea temperatures.Tropicalcorals are affected when seatemperature5rise toapoint at which algaedieoff or'bailout'from theircoralhost;thisis knownascoralbleaching.ln

the absence of the algae,corals are no longer able to sustain themselves resulting inthe eventual death01the coral colony(see Wilkinson,2000).Fordeep-seacoralsthat requirestable temperatures within specific ranges,even a slightchange in temperature at depth could pose a serious threat.Ithas been shown that such changes have begun and temperatures have risen at depths down to 700 m (Barnettetal.,2005).Tolerances of deep-sea coralstosuch temperature fluctuations are unknown

Ocean acidification is another threat to calcareous marineorganisms such as corals (Guinotteetal.,2006;Turleyetal.,2007).ltistriggeredbytherelease ollossil luel by- products (e.g.CO"SO"andNO,) and volcanic ash (CO,)intothe atmosphere,which bond with water particles to lorm carbonic acid (H,C03),sulphuric acid (H,S04) and nitric acid (H,N03).Theseacids eventually precipitate out intotheoceancausing pHlevelsto decreaseand acidity to increase. Ocean acidity has already increasedapproximately 30%since theIndustrialRevolution asa result of anthropogenic carbon released into the atmosphere (Caldeira&Wickett, 2003;Feelyetal.,2008).Adecreasein pH causes calcareous skeletons to erode and weaken,and as aresultcom promisestheintegrityof thesestructures(Feelyetal.,2004;Orretal.,2005;Hall-Spenceretal.,2008)

compostionbydecreasingtheoccurrenceofcalcareousorganismslikescleractinian corals andincreasingsea-grass production (Hall-Spenser et ai.,2008).IIocean acidificationcontinuestorisethelong-termconsequences;though unknown,arelikelyto amplifyphysiologicalstress's on deep-sea corals with similareffectsas seen in shallow

Submarine cables areusedfortelecommunicationpurposes and are spread worldwide

athreat todeep-sea corals because off the direct damage caused duringcablelaying operations.However,fewpeer-reviewed scientificsludieshavebeenwrittento determine the degreeof impactthesecables have onthedeep-sea.One studyhas shownthatsubmarine cableshavea rninoreffect on deep-sea organismsand can provide a suitable hard substrateforattachmentforsome species such as sea anemones and soft corals(Koganetal.,2006).Most old cables are heavily colonized by sessile invertebrates and may be considered positive (Duncan, 1877;Wilson,1979) Otherstudies areneeded todeterminehowsubmarine cablesimpactother species of

1.2.2.6Aquaculture

Newfoundlandaquaculture operations use open-pen systems and areprimarily basedin Nortre Dame Bay,Bayd'EspoirandFortuneBay(FisheriesandAquaculture NewfoundlandandLabrador,2009).Theaquaculture industry is expandingin NewfoundlandandLabrador Region(e.g. salmon,shellfish) and maybeapotential

Newfoundland fiords(Haedrich& Ga9non,1991).Aquaculture operationscanimpact benthicmarineenvironmentsinseveral ways, fromthepollutionreleasedintothe sea as organic waste (Grant &Briggs,1998; Ackefors &Enell,1990;Hargraveet aI.,1993),to the antifoulingagentsusedonthecages which canleach toxicchemicalsintothe sea (Katranitsasetal.,2003).Accumulation of organic waste (e.g. fecalwaste,and unconsumedmedicatedfeedpellets)directlybelow thepencan ereate anoxic

conditions,and asa result reduce species diversity and biomass of benthic macrofauna (Ritzetal.,1989;Weston,1990).Unconsumedmedicated feed pellets can be hazardousas well when consumed by other benthic organisms (Grant&Briggs,1998) Aquaculture in this region impacts primarilyshallowcoastalwaters;thereforeitisonly considered a theatto soft coralsthatare foundincloseproximitytotheaquaculture

Historically,deep-sea coral distributions in thenorthwestAtlanticweredocumentedin the 1800sat the time oftheBlake (Agassiz,1888),Challenger(Moseley, 1881), Albatross(Verrill,188S),andPrinceAlbertofMonaco(Jourdan,189S) expeditions. The Blakeexpedition surveyed thenortheast UnitedStates,while the Challengerand Albatross expeditions surveyed as far north as the Scotian Shelf (Verrill,188S;Agassiz, 1888).Onlythe PrinceAlbert of Monaco expedition (1887) sampledoff Newfoundland (Jourdan,18gS).Duringthisexpeditionbenthicsamplesweretakenat twolocationson

documented at a depth of 1267 m: Caryophyllia communis,Vaughanella margaritata (Jourdan,188S),F.a/abastrum,Anthomastusagaricus(=A.grandifIorus?),Acanella normani(=A. arbuscula?).At Station 162(46°S0'6"N,Soo11'4S"W)onescleractinian species,Desmophyllumdianthus,wasrecordedatadepthofat1SS m(Jourdan,189S) Refer to Figure 1.1 for map illustrating bathymetric features

OntheScotian Shelf offBanquereau Banksignificant coral concentrationswerenoted byCaptain Collins(1884),a prominent fishing skipper.Hewould name the area'The

Inthe early 1900s,the GodthaabExpedition(Kramp,1942)documentedalcyonarians, antipatharians,pennatulaceans,and scleractiniansin the LabradorSea-BaffinBasin area from54°00'N- 79°00'N.Most species were documented off southwest Greenland,withsomespeciesfromBaffin Bay(e.g.sea pens and soft corals),andone soft coral was documented in the vicinity ofHarrison Bank,offcentralLabrador(see Kamp,1932)

Pax(1932)mappedfive occurrences ofStauropathesarclica(;Bathypathes);onesouth

During the 1950s,corals were documented at severallocalitieson theGrand Banksbya groupof Russianscientists (Litvin&Rvachev, 1963;Nesis,1963a;Nesis,1963b).The firststudy generated maps of seabedtopographyand substrates ofthe Newfoundland- Labrador fishingareas (Litvin&Rvachev,1963).Resultsmapped corals attheStone

FlemishCap,and southwestBeothuckKnoll (seeLitvin&Rvachev,1963)

Thesecond study ofthe Russianexpeditionmappedthe bathyal amphiboreal fauna of theNewfoundland-Labradorfishing area (Nesis,1963a).Amphiboreal fauna are defined asspeciesfound in the Pacificand Atlantic borealregionsbutnot intheArctic.Three deep-seacoral species were documented along the continental sheIf break off NewfoundlandandLabrador,with otherrecordsfromthe FlemishCap.Threespecies weredocumented;twolargegorgonians(i.e.Paragorgiaarborea,Primnoa resedaeformis),and onelargesea pen (i.e.Halipterisfinmarchica).Paragorgiaarborea weremappedoffHawkeSaddle,noseofthe Grand Bank, FlemishCap,andmouthof

Channel and on the southwest part ofFlemishCap.Halipteris finmarchica were documented onFlemishCap,and on the southwest GrandBanksnearDesbarres Canyon (seeNesis,1963a).Samples consisted ofby-catchfrom commercial trawls (n=539),Sigsbitrawls (n=99) and bottom grabs (n=164)fromAttantic Canada (Nesis,

ThefinalstudyoftheRussianexpedition(Nesis,1963b)produced mapsof the benthos oftheNewfoundland-Labradorfishingarea.Paragorgiaarborea andP.resedaeformis weredocumentedbetween300-500monthenorthernpartof"GrandNewfoundland Bank"andtheadjacent shelfbreak and slope. Many species of pennatulaceans(i.e AnthoptilumgrandifiorumVerrill,187g;PennatulaaculeataDanieIssen,1860;andP grandisEhrenberg,1834) and one scleractinian cup coral (i.e.F.alabastrum)were documentedbetween 250-500m inLaurentianChannel. Sea pens (i.e.Halipteris finmarchica[=Pavonaria]Sars,1851;andA.grandifiorum),gorgonians(i.e.P.arborea, K.ornata[=Ceratoisis],andR.gracilis),cerianthipatharians(i.e.S.arclica [=Bathypathes]),andhexacorals(i.e.F.a/abastrum)weredocumented between 300-350 montheFlemishCap, break and slope (Nesis,1963b)

In more recentyearsTendal(1992) mapped new and existingrecordsofP.arboreain the NorthAtlantic.Two newrecords were mapped on continental edge of GrandBank, 14on the Scotian Shelf,and12alongthecoast ofNorway.Existingdistributions were mappedoff the southwestern (n=7) and southern (n=4)portionsofGreenland,aswellas alongtheReykjanesRidge(n=2),Faeroelslands(n=5),andNorway(n=1;Tendal, 1992).Overa decadelater, Tendal(2004)mappedthedistributionofStauropathes arctica(=Bathypathes),anantipathariancoralfoundoffsouthwest Greenland adjacent tosoutheastBaffin Island,Canada.Intotal,hemapped 18 knownoccurrences; off

InCanada,advocacy for deep-sea coral protection began in the 1990s in southern Nova Scotiawhentwolocallonglinefishermen,Sanford Atwood andDerekJones,rallied to protect corals in an area known as'Hell Hole',locatedbetweenBrownsBank,in CanadianwatersandGeorgesBank,in United States waters (Breeze et aI.,1997;Lees, 2002).Itwas known as an excellentfishingarea withlarge underwater'trees'and

'strawberry fields',later identified as gorgoniansandsoft corals.Together,thetwo

fishermenformedtheCanadian OceanHabitat ProtectionSociety,thefirstnon-profit organizationdedicatedtoconservingcoralsinCanada.Subsequently,they joined a teamof scientists equipped with aROV,leading theresearch team tothelocation of the first documentedcoral gardensinCanada (Lee,2002)

Breezeetal.(1997) documented local fishers'knowledge on coraI occurrences on the Scotian Shelf, along with distribution information from museum and scientific collections This informationwas compiledintothe first map of coral distributions in Canada.Majority ofcoraloccurencesweremappedalongtheedgeandslopeoftheScotianShelf extendingfromGeorgesBankto theLaurentianChannel

Macisaac et al. (2001) mapped the approximatelocationsof coral by-catch from opportunistic benthic surveys and experimental sites,as well as data collected from groundfish trawl surveys.Datawas gathered primarily from theSeotianShelfwith sporadic occurrencesmappedon the edge and slope ofthe LabradorShelf(i.e.Saglek Bank,OkakBank),Funk Island Bank,andonthenoseofGrandBank

Gass (2002) mapped coral distributionsin thenorthwest Atlantic usingdatafrom;DFO research surveys,fisheries observers andLocal Ecological Knowledge(LEK) fromNova Scotia andNewfoundlandandLabradorfishermen.Themajority ofherfindings were fromtheScotian Shelf,with significant contributions fromNewfoundlandandLabrador

region(Gass&Willison,2005).Coral occurrences were mapped along theedge and slope of the continentalshelf extending fromDavisStraitolfsoutheastBalfinlsland, alongtheLabradorShelf(i.e.SaglekBank,OkakBank,MakkovikBank,HamiltonBank), Northeast NewfoundlandShelf (i.e.BelleIsleBank,OrphanSpur,Tobin'sPoint),Grand Banks (i.e.nose,tail,southwestGrandBanks,Halibut andHaddockChannels),and Scotian Shelf (e.g.StoneFence,The Gully,MisaineHoles,NortheastChannel,Jordan Basin).Wareham andEdinger (2007) used a similar methodology.However,a more systematic approach was used andmorecompletefisheriesobserver data was available,resultingin a greaternumberof samples callected and usedin thisthesis(see Wareham&Edinger,2007;Wareham,2009a)

Mortensenet al.(2006) mappedthe distributionof deep-water coralsinAtlanticCanada usingoverlapping data sources similartoGass and Willison (2005) but added;video surveys (seeMortensenetal.,2006),distributiondatafromprevious studies (Gass, 2002),datafromliteraturesources (i.e.Zibrowius,1980;Kramp,1942;Madsen,1994) andmuseumcollections (e.g.Smithsonian,Atlantic ReferenceCentre,NovaScotia NaturalHistory Museums).The majorityof occurrences weremappedon the Scotian ShelfwithsporadicoccurrencesfromNewfoundland,Labrador,andsoutheastBalfin

Deep-seacoralsarelong-lived,slow-growinganimals(Roarketal.,2006).Thelargest speciesresemble underwatertreesthatcan reachheightsof severalmetres(Mortensen

&Buhl-Mortensen, 2005).Fish utilize largecorals as wellas othercoralhabitatsas

feeding areas, restareas, juvenilefish habitats,andrefuges frompredation (Auster, 2005;Costelloetal.,2005).lnNewfoundlandandLabrador,deep-seacorals are

primarilyfoundin areas of steep bathymetric relief primarily ontheedge and slope of the continental shelf at depths>200m (Nesis,1963ab;Gass&Willison,2005;Mortensenet aI.,2005;Wareham &Edinger,2007;Wareham,2009a). A suite of environmental variablesinfluencecoraldistributions:temperature,strongcurrents,and suitable substrates (Nesis,1963b;Cinbergetal.,1981;Bryan& Metaxas,2006;Mortensen et al.,2006).Manyquestionsstiliremainunansweredconcerningtherelative importance of

Deep-seacorals are ecologically-important because theycreate habitats for other marineorganisms as large individuals oraslargeconcentration5(Austeret aI.,2005;

Buhl-Mortensen& Mortensen,2005; Auster,2007;Etnoyer& Warrenchuk,2007;Moore etal.,2008;Bakeretal.,2008).Whiletherearenumerousthreatstodeep-seacorals, theimpactof bottom fishing gears;particularly trawling,isconsidered to be the greatest threat(Probert,1997;Watling&Norse,1998;Fossaetal.,2001;HaII-Spenseret aI.,

2005;Stone,2006).However,changes in oceanic systems from global warming and ocean acidification are now being documented and potentiallyhavesignificant consequencestocoralsandothermarinelife(Feelyetal.,2004;Orretal.,2005;Hall-

Explorationsconductedinthe late 1800's provided reports on deep-seacoral distributionsinthe northwest Atlantic.Observation advQcacyby local fishersfromNova Scotiaprovidedthe impetus for scientific studies of corals (LeeS,2002),leadingtowell- studied distributions on the Scotian Shelf(Breezeetal.,1997;Maclsaacetal.,2001;

Gass&Wiliison, 2005).A significant outcome ofthis researchhasbeenthe establishmentofthreeimportantareasfordeep-seacoralprotection;The Gully Marine ProlecledArea(DFO,2004a),NorlheastChanneICora/ConservationArea (DFO,

2002),andLophelia[Stone Fence]CoraIConservalionArea(DFO,2004b).These protectionzones willbediscussedin moredetailin Chapter 4

In thecase ofNewfoundlandandLabradorregion,distributions0fdeep-sea coralshave beendocumentedbyearlyexploratoryexpeditions(1800's),andby preceding studies (1963-2006).The nexttwo chapters present geographic and bathymetric distributions (2003-2007) ofdeep-seacorals withinthenorthwest AtlanticfocusingonNewfoundland,

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