Deep-Sea Fishes Biology, Diversity, Ecology and Fisheries Thetechnologicaladvancesofthelasttwentyyearshavebroughthugeadvancesinour understandingofthedeepseaandofthespeciesinhabitingthiselusiveandfascinating environment.Synthesisingtheverylatestresearchanddiscoveries,thisisacomprehensive andmuch-neededaccountofdeep-seafishes.Priedeexaminesallaspectsofthisincredibly diversegroupofanimals,reviewingalmost3,500speciesandcoveringdeep-seafishevolution, physiologyandecologyaswellaschartingthehistoryoftheirdiscoveryfromtheeighteenth centurytothepresentday.Providingaglobalaccountofbothpelagicanddemersalspecies, thebookultimatelyconsiderstheeffectofthegrowingdeep-seafishingindustryon sustainability. Copiouslyillustratedwithexplanationsofthedeep-seaenvironment,drawingsoffishesand informationonhowtheyadapttothedeep,thisisanessentialresourceforbiologists, conservationists,fisherymanagersandanyoneinterestedinmarineevolutionandnatural history. ImantsG.PriedeisProfessorEmeritusatOceanlab,afieldresearchstationoftheInstituteof BiologicalandEnvironmentalSciences,UniversityofAberdeen.Hehasover40yearsof experiencestudyingdeep-seafish,fromlongliningandtrawlingontheRoyalResearchShip Challengerinthe1970stoparticipatinginandleadingexpeditionstothePacificOcean, AtlanticOcean,MediterraneanSeaandtheMid-AtlanticRidge.ThefounderofOceanlab,he alsopioneeredsatellitetrackingofsharksinthe1980sandistherecipientofthe2011 BevertonMedaloftheFishSocietyoftheBritishIsles. Deep-Sea Fishes Biology, Diversity, Ecology and Fisheries IMANTS G. PRIEDE ProfessorEmeritus,Oceanlab,UniversityofAberdeen, Scotland,UK UniversityPrintingHouse,CambridgeCB28BS,UnitedKingdom OneLibertyPlaza,20thFloor,NewYork,NY10006,USA 477WilliamstownRoad,PortMelbourne,VIC3207,Australia 4843/24,2ndFloor,AnsariRoad,Daryaganj,Delhi-110002,India 79AnsonRoad,#06-04/06,Singapore079906 CambridgeUniversityPressispartoftheUniversityofCambridge. ItfurtherstheUniversity’smissionbydisseminatingknowledgeinthepursuitof education,learning,andresearchatthehighestinternationallevelsofexcellence. www.cambridge.org Informationonthistitle:www.cambridge.org/9781107083820 DOI:10.1017/9781316018330 ©ImantsG.Priede2017 Thispublicationisincopyright.Subjecttostatutoryexception andtotheprovisionsofrelevantcollectivelicensingagreements, noreproductionofanypartmaytakeplacewithoutthewritten permissionofCambridgeUniversityPress. Firstpublished2017 PrintedintheUnitedKingdombyTJInternationalLtd.PadstowCornwall AcataloguerecordforthispublicationisavailablefromtheBritishLibrary. LibraryofCongressCataloging-in-PublicationData Names:Priede,I.G.,author. Title:Deep-seafishes:biology,diversity,ecologyandfisheries/ImantsG.Priede,professoremeritus,Oceanlab, UniversityofAberdeen,Scotland,UK. Description:Cambridge,UnitedKingdom;NewYork,NY:CambridgeUniversityPress,2017.|Includes bibliographicalreferencesandindex. Identifiers:LCCN2017006037|ISBN9781107083820(Hardback:alk.paper) Subjects:LCSH:Deep-seafishes.|Deep-seabiology.|Deep-seaecology. Classification:LCCQL620.P752017|DDC597/.63–dc23LCrecordavailableathttps://lccn.loc.gov/2017006037 ISBN978-1-107-08382-0Hardback CambridgeUniversityPresshasnoresponsibilityforthepersistenceoraccuracy ofURLsforexternalorthird-partyinternetwebsitesreferredtointhispublication anddoesnotguaranteethatanycontentonsuchwebsitesis,orwillremain, accurateorappropriate. To my wife, Maria. CONTENTS Preface pageix 1.7.2 MethodsofFishCapture 51 1.7.3 ImagingSystems 61 1 Introduction 1 1.7.4 TaggingandTrackingofFishes 65 1.1 DefinitionofDeep-SeaFishes 1 1.2 TheStructureoftheWorld’sOcean 2 Colonisation of the Deep Sea by Fishes 68 Basins 2 2.1 Introduction 68 1.2.1 OceanDepthZones 2 2.2 Present-DayDepthDistributionofFishes 68 1.2.2 OceanBasinFormation,Slopes, 2.3 TheHistoryofDeep-SeaFishes 72 Plains,Ridges,Islandsand 2.4 OriginsoftheModernDeep-SeaFishFauna 76 Seamounts 6 2.4.1 Agnatha 76 1.2.3 TheBathyalSlopes 2.4.2 Chondrichthyes 76 (200–3000mDepth) 11 2.4.3 Actinopterygii 79 1.2.4 HydrothermalVentsand 2.4.4 Sarcopterygii 85 ColdSeeps 12 2.5 AncientandSecondaryDeep-SeaFishes 85 1.2.5 TheAbyss(3000–6000mDepth) 13 1.2.6 TheHadalZone(>6000mDepth) 15 3 Adaptations to the Deep Sea 87 1.3 HistoryoftheOceanBasins 16 3.1 Introduction 87 1.4 TheDeep-SeaEnvironment 21 3.2 EnergyMetabolism 87 1.4.1 DeepWaterFormation,Circulation 3.3 EffectsofPressure 90 andOxygenationoftheDeep 21 3.4 SizeandShape 91 1.4.2 SurfaceOceanCirculationandGyres 27 3.5 Buoyancy 93 1.4.3 OceanBiomes 27 3.5.1 TheSwimBladderandGasGland 93 1.5 FoodSupplytotheDeepandthe 3.5.2 LipidBuoyancy 100 DistributionofBiomass 30 3.5.3 WeightReduction 101 1.5.1 ExportofParticulateOrganic 3.6 MuscleAdaptations 102 MatterfromtheSurface 30 3.7 GillArea 103 1.5.2 FoodFalls 31 3.8 HeartandCirculatorySystem 104 1.5.3 DistributionofBiomassin 3.9 TheArtofSlowSwimmingandthe theDeepSea 31 OptimumSpeedFallacy 105 1.6 HistoryofDiscoveryofDeep-SeaFishes 35 3.10 FeedingandGrowth 108 1.6.1 FromtheBeginningsofIchthyology 3.11 LifeHistoryStrategies 114 totheFirstDeep-SeaDiscoveries, 3.12 Excretion 118 1554–1860 35 3.13 SensoryandCommunicationSystems 119 1.6.2 GreatVoyagesofDiscovery, 3.13.1 DiversityofBrainMorphology 119 1860–1914 38 3.13.2 Vision 122 1.6.3 TotheFirstMannedDescentsinto 3.13.3 CamouflageintheDeep 125 theDeep,1914–1945 41 3.13.4 Bioluminescence 127 1.6.4 TheGreatTwentieth-Century 3.13.5 TheOlfactoryandGustatory ScientificExpansionfrom1945 42 Systems 133 1.7 CollectionandMethods 49 3.13.6 UnderwaterSoundandthe 1.7.1 AccessingtheDeepSea 49 OctavoLateralSystem 133 viii Contents 3.14 TimeintheDeepSea 135 4.3.25 OrderTetraodontiformes 299 3.15 DiseasesandtheImmuneSystem 136 4.3.26 OrderLophiiformes 301 3.16 TheMaximumDepthLimitforFishes 138 4.4 ClassSarcopterygii 314 4.4.1 OrderCoelacanthiformes 314 4 Systematic Description of Deep-Sea Fishes 139 5 The Deep-Sea Fish Faunas 317 4.1 ClassMyxine 139 5.1 MesopelagicandBathypelagicFish 4.1.1 OrderMyxiniformes 139 Fauna 317 4.1.2 OrderPetromyzontiformes 141 5.1.1 ArcticOcean 319 4.2 ClassChondrichthyes 142 5.1.2 PacificOcean 319 4.2.1 OrderChimaeriformes 142 5.1.3 IndianOcean 333 4.2.2 OrderHexanchiformes 144 5.1.4 AtlanticOcean 334 4.2.3 OrderPristiophoriformes 144 5.1.5 TheAntarcticOcean 335 4.2.4 OrderSquatiniformes 145 5.2 BathyalDemersalFishes 4.2.5 OrderSqualiformes 145 (200–3000mDepth) 335 4.2.6 OrderLamniformes 149 5.3 AbyssalDemersalSpecies 4.2.7 OrderCarcharhiniformes 150 (3000–6000mDepth) 347 4.2.8 OrderTorpediformes 152 5.4 FishesoftheHadalZone 4.2.9 OrderPristiformes 152 (>6000mDepth) 356 4.2.10 OrderMyliobatiformes 152 5.5 FishesofHydrothermalVentsand 4.2.11 OrderRajiformes 153 ColdSeeps 358 4.3 ClassActinopterygii 155 5.6 TheCircumglobalDeep-SeaFishFauna 362 4.3.1 OrderAlbuliformes 155 6 Deep-Sea Fisheries and Conservation 363 4.3.2 OrderNotacanthiformes 156 4.3.3 OrderAnguilliformes 158 6.1 GlobalTrendsinDeepFishing 363 4.3.4 OrderAlepocephaliformes 166 6.2 TypesofFishing 364 4.3.5 OrderArgentiniformes 171 6.2.1 BaitedTrapsandLines 364 4.3.6 OrderStomiiformes 175 6.2.2 StaticNets 364 4.3.7 OrderAteleopodiformes 186 6.2.3 PelagicTrawling 364 4.3.8 OrderAulopiformes 187 6.2.4 BottomTrawling 365 4.3.9 OrderMyctophiformes 196 6.3 GlobalTrendsinFishCatch 365 4.3.10 OrderLampriformes 203 6.4 CompositionoftheWorldDeep-Sea 4.3.11 OrderPolymixiiformes 207 FishCatch 367 4.3.12 OrderZeiformes 208 6.4.1 TheTopTwentyDeep-SeaSpecies 384 4.3.13 OrderStylephoriformes 210 6.4.2 OtherSpecies 391 4.3.14 OrderGadiformes 210 6.5 OverviewoftheFishingTrends 400 4.3.15 OrderOphidiiformes 228 6.6 TheImpactsofDeep-SeaFishing 400 4.3.16 OrderBatrachoidiformes 238 6.7 Conclusions 403 4.3.17 OrderGobiesociformes 238 Epilogue 404 4.3.18 OrderStephanoberyciformes 239 4.3.19 OrderBeryciformes 241 4.3.20 OrderCetomimiformes 245 Appendix 406 4.3.21 OrderSyngnathiformes 248 References 415 4.3.22 OrderScorpaeniformes 249 Index 470 4.3.23 OrderPerciformes 264 Colourplatescanbefoundbetweenpages252and 4.3.24 OrderPleuronectiformes 294 253.