Fishes of the World Fishes of the World Fifth Edition Joseph S. Nelson Terry C. Grande Mark V. H. Wilson Coverimage:MarkV.H.Wilson Coverdesign:Wiley Thisbookisprintedonacid-freepaper. Copyright©2016byJohnWiley&Sons,Inc.Allrightsreserved. PublishedbyJohnWiley&Sons,Inc.,Hoboken,NewJersey. PublishedsimultaneouslyinCanada. Nopartofthispublicationmaybereproduced,storedinaretrievalsystem,ortransmittedinany formorbyanymeans,electronic,mechanical,photocopying,recording,scanning,orotherwise, exceptaspermittedunderSection107or108ofthe1976UnitedStatesCopyrightAct,without eitherthepriorwrittenpermissionofthePublisher,orauthorizationthroughpaymentofthe appropriateper-copyfeetotheCopyrightClearanceCenter,222RosewoodDrive,Danvers,MA 01923,(978)750-8400,fax(978)646-8600,oronthewebatwww.copyright.com.Requeststothe PublisherforpermissionshouldbeaddressedtothePermissionsDepartment,JohnWiley& Sons,Inc.,111RiverStreet,Hoboken,NJ07030,(201)748-6011,fax(201)748-6008,oronlineat www.wiley.com/go/permissions. 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Classification:LCCQL618.N42016|DDC597.01/2–dc23LCrecordavailableat http://lccn.loc.gov/2015037522 PrintedintheUnitedStatesofAmerica 10 9 8 7 6 5 4 3 2 1 Contents MEMORIESOFJOENELSON xxix FOREWORD xxxiii PREFACE xxxvii ACKNOWLEDGMENTS xxxix INTRODUCTION 1 TaxonomicDiversity 3 ImportancetoPeople 4 SystematicsandClassification 4 AnatomicalTerminology 7 DistributionandBiogeography 8 HumanImpacts 11 PHYLUMCHORDATA 13 SUBPHYLUMUROCHORDATA(Tunicata:thetunicates) 15 ClassASCIDIACEA(ascidians), 15 ClassTHALIACEA(salps), 15 OrderPYROSOMIDA, 15 OrderDOLIOLIDA, 15 OrderSALPIDA, 15 ClassAPPENDICULARIA, 15 SUBPHYLUMCEPHALOCHORDATA(Acrania,inpart) 16 OrderAMPHIOXIFORMES(lancelets), 16 FamilyBRANCHIOSTOMATIDAE, 16 FamilyEPIGONICHTHYIDAE, 16 v vi Contents †SUBPHYLUMCONODONTOPHORIDA(conodonts) 17 †ClassCONODONTA, 17 SUBPHYLUMCRANIATA 18 INFRAPHYLUMMYXINOMORPHI 19 ClassMYXINI, 20 OrderMYXINIFORMES(1)—hagfishes, 20 FamilyMYXINIDAE(1)—hagfishes, 20 INFRAPHYLUMVERTEBRATA(vertebrates) 22 SUPERCLASSPETROMYZONTOMORPHI 23 ClassPETROMYZONTIDA, 23 OrderPETROMYZONTIFORMES(2)—lampreys, 23 †FamilyMAYOMYZONTIDAE, 24 FamilyPETROMYZONTIDAE(2)—northernlampreys, 24 FamilyGEOTRIIDAE(3)—southernlampreys, 25 FamilyMORDACIIDAE(4)—southerntopeyedlampreys, 26 †SUPERCLASSPTERASPIDOMORPHI 26 †ClassPTERASPIDOMORPHA, 26 †SubclassASTRASPIDA, 27 †OrderASTRASPIDIFORMES, 27 †SubclassARANDASPIDA, 27 †OrderARANDASPIDIFORMES, 27 †SubclassHETEROSTRACI, 28 †CARDIPELTIDA, 28 †CORVASPIDIDA, 28 †LEPIDASPIDIDA, 28 †TESSERASPIDIDA, 28 †TRAQUAIRASPIDIFORMES, 28 †TOLYPELEPIDIDA, 28 †OrderCYATHASPIDIFORMES, 28 †FamilyAMPHIASPIDIDAE, 28 †FamilyCYATHASPIDIDAE, 29 †OrderPTERASPIDIFORMES, 29 †FamilyANCHIPTERASPIDIDAE, 29 †FamilyPROTASPIDIDAE, 29 †FamilyPROTOPTERASPIDIDAE, 29 †FamilyPSAMMOSTEIDAE, 29 †FamilyPTERASPIDIDAE, 29 †SUPERCLASSANASPIDOMORPHI 29 †ClassANASPIDA, 29 †OrderANASPIDIFORMES, 29 †SUPERCLASSTHELODONTOMORPHI 30 †ClassTHELODONTI, 30 †OrderARCHIPELEPIDIFORMES, 31 †OrderFURCACAUDIFORMES(fork-tailedthelodonts), 31 †OrderTHELODONTIFORMES, 31 Contents vii †SUPERCLASSOSTEOSTRACOMORPHI 32 †ClassCEPHALASPIDOMORPHI, 33 †OrderCEPHALASPIDIFORMES, 33 †OrderGALEASPIDIFORMES, 34 †OrderPITURIASPIDIFORMES, 34 SUPERCLASSGNATHOSTOMATA(jawedvertebrates) 34 †GradePLACODERMIOMORPHI 36 †ClassPLACODERMI, 36 †OrderPSEUDOPETALICHTHYIFORMES, 37 †OrderACANTHOTHORACIFORMES, 37 †OrderRHENANIFORMES, 37 †OrderANTIARCHIFORMES(antiarchs), 37 †FamilyCHUCHINOLEPIDIDAE, 38 †FamilyYUNNANOLEPIDIDAE, 38 †FamilySINOLEPIDAE, 38 †FamilyMICROBRACHIIDAE, 38 †FamilyBOTHRIOLEPIDIDAE, 38 †FamilyASTEROLEPIDIDAE, 38 †FamilyGERDALEPIDIDAE, 39 †OrderPETALICHTHYIFORMES, 39 †OrderPTYCTODONTIFORMES, 39 †OrderARTHRODIRIFORMES(arthrodires), 39 EUGNATHOSTOMATA, 40 GRADECHONDRICHTHYOMORPHI 40 ClassCHONDRICHTHYES—cartilaginousfishes, 40 †FamilyPROTODONTIDAE, 43 †FamilyKATHEMACANTHIDAE, 43 †OrderPOLYMEROLEPIDIFORMES, 44 †OrderOMALODONTIFORMES, 44 †OrderANTARCTILAMNIFORMES, 44 †FamilyBANDRINGIDAE, 45 †OrderPHOEBODONTIFORMES, 45 †FamilyPHOEBODONTIDAE, 45 †FamilyJALODONTIDAE, 45 †SuperorderCLADOSELACHIMORPHA, 46 †OrderCLADOSELACHIFORMES, 46 †FamilyCLADOSELACHIDAE, 46 †OrderSYMMORIIFORMES, 46 †FamilySYMMORIIDAE, 46 †FamilyFALCATIDAE, 46 †SuperorderCTENACANTHIMORPHA, 46 †OrderCTENACANTHIFORMES, 46 †FamilyCTENACANTHIDAE, 47 †OrderSQUATINACTIFORMES, 47 †FamilySQUATINACTIDAE, 47 †SuperorderXENACANTHIMORPHA, 47 †OrderBRANSONELLIFORMES, 47 viii Contents †OrderXENACANTHIFORMES, 47 †FamilyDIPLODOSELACHIDAE, 47 †FamilyXENACANTHIDAE, 47 SubclassHOLOCEPHALI, 48 †OrderINIOPTERYGIFORMES, 49 †OrderORODONTIFORMES, 49 †OrderEUGENEODONTIFORMES, 50 †OrderPETALODONTIFORMES, 50 †OrderDEBEERIIFORMES, 50 †OrderHELODONTIFORMES, 50 SuperorderHOLOCEPHALIMORPHA, 50 †OrderPSAMMODONTIFORMES, 50 †OrderCOPODONTIFORMES, 50 †OrderSQUALORAJIFORMES, 51 †OrderCHONDRENCHELYIFORMES, 51 †OrderMENASPIFORMES, 51 †OrderCOCHLIODONTIFORMES, 51 OrderCHIMAERIFORMES(3)—chimaeras, 51 FamilyCALLORHINCHIDAE(5)—plownosechimaeras, 52 FamilyRHINOCHIMAERIDAE(6)—longnosechimaeras, 52 FamilyCHIMAERIDAE(7)—shortnosechimaerasorratfishes, 53 SubclassEUSELACHII(sharks,rays,andrelatedfossils), 53 †OrderPROTACRODONTIFORMES, 55 †InfraclassHYBODONTA, 55 †OrderHYBODONTIFORMES, 55 †FamilyHYBODONTIDAE, 56 †FamilyTRISTYCHIIDAE, 56 †FamilyDISTOBATIDAE, 56 †FamilyACRODONTIDAE, 56 †FamilyPOLYACRODONTIDAE, 56 †FamilyLONCHIDIIDAE, 56 †FamilySTEINBACHODONTIDAE, 56 †FamilyPSEUDODALATIIDAE, 56 †FamilyPTYCHODONTIDAE, 56 †FamilyHOMALODONTIDAE, 56 InfraclassELASMOBRANCHII, 56 DivisionSELACHII—sharks, 58 SuperorderGALEOMORPHI, 58 †OrderSYNECHODONTIFORMES, 59 †FamilyPALAEOSPINACIDAE, 59 OrderHETERODONTIFORMES(4)—bullheadsharks, 59 FamilyHETERODONTIDAE(8)—bullheadsharks, 59 OrderORECTOLOBIFORMES(5)—carpetsharks, 59 FamilyPARASCYLLIIDAE(9)—collaredcarpetsharks, 60 FamilyBRACHAELURIDAE(10)—blindsharks, 60 FamilyORECTOLOBIDAE(11)—wobbegongs, 61
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