Main Fishes of the world, 5th Edition.

Fishes of the world, 5th Edition.

, ,
Year: 2016
Edition: 5
Publisher: Wiley
Language: english
Pages: 752
ISBN 13: 9781118342336
File: AZW3 , 16.30 MB
Download (azw3, 16.30 MB)

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Table of Contents


Title Page

Copyright

Memories of Joe Nelson

Foreword

Preface

Acknowledgments

Introduction Taxonomic Diversity

Importance to People

Systematics and Classification

Anatomical Terminology

Distribution and Biogeography

Human Impacts





Phylum Chordata SUBPHYLUM UROCHORDATA (Tunicata: the tunicates) Class ASCIDIACEA (ascidians)

Class THALIACEA (salps) Order PYROSOMIDA

Order DOLIOLIDA (Cyclomyaria)

Order SALPIDA (Hemimyaria)





Class APPENDICULARIA (Larvacea)





SUBPHYLUM CEPHALOCHORDATA (Acrania, in part) Order AMPHIOXIFORMES (lancelets) Family BRANCHIOSTOMATIDAE

Family EPIGONICHTHYIDAE (Asymmetrontidae)





† SUBPHYLUM CONODONTOPHORIDA (conodonts) †Class CONODONTA





SUBPHYLUM CRANIATA

INFRAPHYLUM MYXINOMORPHI Class MYXINI Order MYXINIFORMES (Hyperotreti) (1)—hagfishes Family MYXINIDAE (1)—hagfishes





INFRAPHYLUM VERTEBRATA (vertebrates)

SUPERCLASS PETROMYZONTOMORPHI Class PETROMYZONTIDA Order PETROMYZONTIFORMES (Hyperoartii) (2)—lampreys †Family MAYOMYZONTIDAE

Family PETROMYZONTIDAE (2)—northern lampreys

Family GEOTRIIDAE (3)—southern lampreys

Family MORDACIIDAE (4)—southern topeyed lampreys





†SUPERCLASS PTERASPIDOMORPHI †Class PTERASPIDOMORPHA (Diplorhina)

†Subclass ASTRASPIDA †Order ASTRASPIDIFORMES





†Subclass ARANDASPIDA †Order ARANDASPIDIFORMES





†Subclass HETEROSTRACI †CARDIPELTIDA

†CORVASPIDIDA

†LEPIDASPIDIDA

†TESSERASPIDIDA

†TRAQUAIRASPIDIFORMES

†TOLYPELEPIDIDA

†Order CYATHASPIDIFORMES †Family AMPHIASPIDIDAE

†Family CYATHASPIDIDAE





†Order PTERASPIDIFORMES †Family ANCHIPTERASPIDIDAE

†Family PROTASPIDIDAE

†Family PROTOPTERASPIDIDAE

†Family PSAMMOSTEIDAE

†Family PTERASPIDIDAE





†SUPERCLASS ANASPIDOMORPHI †Class ANASPIDA †Order ANASPIDIFORMES (Birkeniae, Birkeniida)





†SUPERCLASS THELODONTOMORPHI †Class THELODONTI †Order ARCHIPELEPIDIFORMES

†Order FURCACAUDIFORMES (fork-tailed thelodonts)

†Order THELODONTIFORMES





†SUPERCLASS OSTEOSTRACOMORPHI †Class CEPHALASPIDOMORPHI (Monorhina) †Order CEPHALASPIDIFORMES (Osteostraci)

†Order GALEASPIDIFORMES

†Order PITURIASPIDIFORMES (Pituriaspida)





SUPERCLASS GNATHOSTOMATA (jawed vertebrates)

†Grade PLACODERMIOMORPHI †Class PLACODERMI †Order PSEUDOPETALICHTHYIFORMES

†Order ACANTHOTHORACIFORMES

†Order RHENANIFORMES

†Order ANTIARCHIFORMES (antiarchs) †Family CHUCHINOLEPIDIDAE

†Family YUNNANOLEPIDIDAE

†Family SINOLEPIDAE

†Family MICROBRACHIIDAE

†Family BOTHRIOLEPIDIDAE

†Family ASTEROLEPIDIDAE

†Family GERDALEPIDIDAE





†Order PETALICHTHYIFORMES

†Order PTYCTODONTIFORMES

†Order ARTHRODIRIFORMES (arthrodires)





EUGNATHOSTOMATA





GRADE CHONDRICHTHYOMORPHI Class CHONDRICHTHYES—cartilaginous fishes †Family PROTODONTIDAE

†Family KATHEMACANTHIDAE

†Order POLYMEROLEPIDIFORMES

†Order OMALODONTIFORMES

†Order ANTARCTILAMNIFORMES †Family BANDRINGIDAE





†Order PHOEBODONTIFORMES †Family PHOEBODONTIDAE

†Family JALODONTIDAE





†Superorder CLADOSELACHIMORPHA †Order CLADOSELACHIFORMES †Family CLADOSELACHIDAE





†Order SYMMORIIFORMES †Family SYMMORIIDAE

†Family FALCATIDAE





†Superorder CTENACANTHIMORPHA †Order CTENACANTHIFORMES †Family CTENACANTHIDAE





†Order SQUATINACTIFORMES †Family SQUATINACTIDAE





†Superorder XENACANTHIMORPHA (Pleuracanthodii) †Order BRANSONELLIFORMES

†Order XENACANTHIFORMES †Family DIPLODOSELACHIDAE

†Family XENACANTHIDAE





Subclass HOLOCEPHALI †Order INIOPTERYGIFORMES

†Order ORODONTIFORMES

†Order EUGENEODONTIFORMES

†Order PETALODONTIFORMES

†Order DEBEERIIFORMES

†Order HELODONTIFORMES





Superorder HOLOCEPHALIMORPHA †Order PSAMMODONTIFORMES

†Order COPODONTIFORMES

†Order SQUALORAJIFORMES

†Order CHONDRENCHELYIFORMES

†Order MENASPIFORMES

†Order COCHLIODONTIFORMES

Order CHIMAERIFORMES (3)—chimaeras Family CALLORHINCHIDAE (Callorhynchidae) (5)—plownose chimaeras

Family RHINOCHIMAERIDAE (6)—longnose chimaeras

Family CHIMAERIDAE (7)—shortnose chimaeras or ratfishes





Subclass EUSELACHII (sharks, rays, and related fossils) †Order PROTACRODONTIFORMES





†Infraclass HYBODONTA (hybodonts) †Order HYBODONTIFORMES †Family HYBODONTIDAE

†Family TRISTYCHIIDAE

†Family DISTOBATIDAE

†Family ACRODONTIDAE

†Family POLYACRODONTIDAE

†Family LONCHIDIIDAE

†Family STEINBACHODONTIDAE

†Family PSEUDODALATIIDAE

†Family PTYCHODONTIDAE

†Family HOMALODONTIDAE





Infraclass ELASMOBRANCHII (= NEOSELACHII)

Division SELACHII—sharks

Superorder GALEOMORPHI †Order SYNECHODONTIFORMES †Family PALAEOSPINACIDAE





Order HETERODONTIFORMES (4)—bullhead sharks Family HETERODONTIDAE (8)—bullhead sharks





Order ORECTOLOBIFORMES (5)—carpet sharks Family PARASCYLLIIDAE (9)—collared carpet sharks

Family BRACHAELURIDAE (10)—blind sharks

Family ORECTOLOBIDAE (11)—wobbegongs

Family HEMISCYLLIIDAE (12)—bamboo sharks

Family GINGLYMOSTOMATIDAE (13)—nurse sharks

Family STEGOSTOMATIDAE (14)—zebra sharks

Family RHINCODONTIDAE (Rhiniodontidae) (15)—whale sharks





Order LAMNIFORMES (6)—mackerel sharks †Family OTODONTIDAE

†Family XIPHODOLAMIIDAE

†Family CARDABIODONTIDAE

†Family CRETOXYRHINIDAE

†Family ARCHAEOLAMNIDAE

†Family PSEUDOSCAPANORHYNCHIDAE

†Family ANACORACIDAE

†Family PSEUDOCORACIDAE

Family MITSUKURINIDAE (16)—goblin sharks

Family ODONTASPIDIDAE (17)—sand tiger sharks

Family PSEUDOCARCHARIIDAE (18)—crocodile sharks

Family ALOPIIDAE (19)—thresher sharks

Family MEGACHASMIDAE (20)—megamouth sharks

Family CETORHINIDAE (21)—basking sharks

Family LAMNIDAE (22)—mackerel sharks





Order CARCHARHINIFORMES (7)—ground sharks Family SCYLIORHINIDAE (23)—cat sharks

Family PROSCYLLIIDAE (24)—finback cat sharks

Family PSEUDOTRIAKIDAE (25)—false cat sharks

Family LEPTOCHARIIDAE (26)—barbeled hound sharks

Family TRIAKIDAE (27)—hound sharks

Family HEMIGALEIDAE (28)—weasel sharks

Family CARCHARHINIDAE (29)—requiem sharks

Family SPHYRNIDAE (30)—hammerhead sharks





Superorder SQUALOMORPHI Series HEXANCHIDA

Order HEXANCHIFORMES (Notidanoidei) (8)—six-gill sharks Family CHLAMYDOSELACHIDAE (31)—frilled sharks

Family HEXANCHIDAE (32)—cow sharks





Series SQUALIDA

Order SQUALIFORMES (9)—dogfish sharks Family CENTROPHORIDAE (33)—gulper sharks

Family ETMOPTERIDAE (34)—lantern sharks

Family SOMNIOSIDAE (35)—sleeper sharks

Family OXYNOTIDAE (36)—rough sharks

Family DALATIIDAE (37)—kitefin sharks

Family SQUALIDAE (38)—dogfish sharks





Series SQUATINIDA

†Order PROTOSPINACIFORMES

Order ECHINORHINIFORMES (10)—bramble sharks Family ECHINORHINIDAE (39)—bramble sharks





Order SQUATINIFORMES (11)—angel sharks Family SQUATINIDAE (40)—angel sharks





Order PRISTIOPHORIFORMES (12)—saw sharks

Family PRISTIOPHORIDAE (41)—saw sharks





Division BATOMORPHI—rays Order TORPEDINIFORMES (13)—electric rays †Family ARCHAEOBATIDAE

Family TORPEDINIDAE (42)—torpedo electric rays

Family NARCINIDAE (43)—numbfishes





Order RAJIFORMES (14)—skates †Family Cyclobatidae

Family RAJIDAE (44)—skates





Order PRISTIFORMES (15)—guitarfishes and sawfishes †Family Sclerorhynchidae

†Family Ptychotrygonidae

Family “RHINOBATIDAE” (45)—guitarfishes

Family RHINIDAE (46)—bowmouth guitarfishes

Family RHYNCHOBATIDAE (47)—wedgefishes

Family PRISTIDAE (48)—sawfishes





Order MYLIOBATIFORMES (16)—stingrays Family PLATYRHINIDAE (49)—thornbacks

Family ZANOBATIDAE (50)—panrays

Family PLESIOBATIDAE (51)—deepwater stingrays

Family UROLOPHIDAE (52)—round stingrays

Family HEXATRYGONIDAE (53)—sixgill stingrays

Family DASYATIDAE (Trygonidae) (54)—whiptail stingrays

Family POTAMOTRYGONIDAE (55)—river stingrays

Family GYMNURIDAE (56)—butterfly rays

Family UROTRYGONIDAE (57)—American round stingrays

Family MYLIOBATIDAE (58)—eagle rays





Grade TELEOSTOMI †Class ACANTHODII—acanthodians †Order CLIMATIIFORMES †Family BROCHOADMONIDAE

†Family CLIMATIIDAE

†Family GYRACANTHIDAE

†Family EUTHACANTHIDAE





†Order DIPLACANTHIFORMES †Family CULMACANTHIDAE

†Family DIPLACANTHIDAE

†Family GLADIOBRANCHIDAE





†Order ISCHNACANTHIFORMES †Family ISCHNACANTHIDAE

†Family PORACANTHODIDAE





†Order ACANTHODIFORMES †Family MESACANTHIDAE

†Family CHEIRACANTHIDAE

†Family ACANTHODIDAE





Class OSTEICHTHYES—bony fishes and tetrapods (= Euteleostomi of Nelson, 2006) †Order DIALIPINIFORMES





Subclass SARCOPTERYGII—lobe-finned fishes and tetrapods

Infraclass ACTINISTIA—coelacanths (Coelacanthida) Order COELACANTHIFORMES (17)—coelacanths †Family MIGUASHAIIDAE

†Family DIPLOCERCIDAE

†Family HADRONECTORIDAE

†Family RHABDODERMATIDAE

†Family LAUGIIDAE

†Family WHITEIIDAE

†Family REBELLATRICIDAE

†Family COELACANTHIDAE

†Family MAWSONIIDAE

Family LATIMERIIDAE (59)—gombessas or coelacanths





Infraclass ONYCHODONTIDA †Order ONYCHODONTIFORMES (Struniiformes) †Family ONYCHODONTIDAE





Infraclass DIPNOMORPHA

†Superorder POROLEPIMORPHA †Order POROLEPIFORMES (Holoptychiiformes) †Family POROLEPIDAE

†Family HOLOPTYCHIIDAE





Superorder DIPNOI (Dipterimorpha, Dipnoiformes of Cloutier and Ahlberg, 1996) †Order DIABOLEPIDIFORMES †Family DIABOLEPIDIDAE





†Order DIPNORHYNCHIFORMES †Family URANOLOPHIDAE

†Family DIPNORHYNCHIDAE

†Family CHIRODIPTERIDAE





†Order Dipteriformes †Family STOMIAHYKIDAE

†Family DIPTERIDAE

†Family CHIRODIPTERIDAE

†Family RHYNCHODIPTERIDAE

†Family PHANEROPLEURIDAE

†Family FLEURANTIIDAE





†Order CTENODONTIFORMES †Family URONEMIDAE

†Family CTENODONTIDAE





Order CERATODONTIFORMES (18)—living lungfishes and their fossil relatives †Family ARGANODONTIDAE

†Family CERATODONTIDAE

†Family ASIATOCERATODONTIDAE

Family NEOCERATODONTIDAE (60)—Australian lungfishes

Family LEPIDOSIRENIDAE (61)—South American lungfishes

Family PROTOPTERIDAE (62)—African lungfishes





†Infraclass RHIZODONTIDA (Rhizodontimorpha) †Order RHIZODONTIFORMES †Family RHIZODONTIDAE





†Infraclass OSTEOLEPIDIDA †Order OSTEOLEPIDIFORMES †Family CANOWINDRIDAE

†Family MEGALICHTHYIDAE

†Family OSTEOLEPIDIDAE

†Family TRISTICHOPTERIDAE

†Family RHIZODOPSIDAE





†Infraclass ELPISTOSTEGALIA †Order ELPISTOSTEGALIFORMES †Family ELPISTOSTEGALIDAE (†PANDERICHTHYIDAE)





Infraclass TETRAPODA—tetrapods

Subclass ACTINOPTERYGII—ray-finned fishes †Family HAPLOLEPIDAE

†Order CHEIROLEPIDIFORMES

†Order PALAEONISCIFORMES

†Order DORYPTERIFORMES †Family DORYPTERIDAE





†Order PLATYSOMIFORMES †Family PLATYSOMIDAE

†Family BOBASATRANIIDAE





†Order TARRASIIFORMES †Family TARRASIIDAE





†Order GUILDAYICHTHYIFORMES †Family GUILDAYICHTHYIDAE





†Order PHANERORHYNCHIFORMES †Family PHANERORHYNCHIDAE





†Order SAURICHTHYIFORMES †Family SAURICHTHYIDAE





†Order REDFIELDIIFORMES †Family REDFIELDIIDAE





†Order PTYCHOLEPIDIFORMES †Family PTYCHOLEPIDIDAE





†Order PHOLIDOPLEURIFORMES †Family PHOLIDOPLEURIDAE





†Order PERLEIDIFORMES †Family CEPHALOXENIDAE

†Family CLEITHROLEPIDIDAE

†Family COLOBODONTIDAE

†Family PERLEIDIDAE

†Family PLATYSIAGIDAE





†Order LUGANOIFORMES †Family LUGANOIDAE





†Order PERLEIDIFORMES





Infraclass CLADISTIA Order POLYPTERIFORMES (Brachiopterygii) (19)—bichirs Family POLYPTERIDAE (63)—bichirs





Infraclass CHONDROSTEI †Order CHONDROSTEIFORMES †Family CHONDROSTEIDAE





Order ACIPENSERIFORMES (20)—paddlefishes and sturgeons †Family PEIPIAOSTEIDAE

Family POLYODONTIDAE (64)—paddlefishes

Family ACIPENSERIDAE (65)—sturgeons





NEOPTERYGII †Order PYCNODONTIFORMES





Infraclass HOLOSTEI (gars, bowfins, and relatives)

Division GINGLYMODI †Order DAPEDIIFORMES †Family DAPEDIIDAE





Order LEPISOSTEIFORMES (21)—gars †Family OBAICHTHYIDAE

Family LEPISOSTEIDAE (66)—gars





†Order SEMIONOTIFORMES †Family SEMIONOTIDAE (Lepidotidae)

†Family CALLIPURBECKIDAE





†Order MACROSEMIIFORMES †Family MACROSEMIIDAE





Division HALECOMORPHI †Order PARASEMIONOTIFORMES

†Order IONOSCOPIFORMES

Order AMIIFORMES (22)—bowfins †Family CATURIDAE

†Family LIODESMIDAE

†Family SINAMIIDAE

Family AMIIDAE (67)—bowfins





Division TELEOSTEOMORPHA

†Subdivision ASPIDORHYNCHEI †Order ASPIDORHYNCHIFORMES †Family ASPIDORHYNCHIDAE





†Order PACHYCORMIFORMES †Family PACHYCORMIDAE





Subdivision TELEOSTEI †Order PHOLIDOPHORIFORMES †Family PHOLIDOPHORIDAE





†Order DORSETICHTHYIFORMES †Family DORSETICHTHYIDAE





†Order LEPTOLEPIDIFORMES †Family LEPTOLEPIDIDAE





†Order CROSSOGNATHIFORMES †Family VARASICHTHYIDAE

†Family CROSSOGNATHIDAE

†Family NOTELOPIDAE

†Family PACHYRHIZODONTIDAE





†Order ICHTHYODECTIFORMES †Family ALLOTHRISSOPIDAE

†Family CLADOCYCLIDAE

†Family SAURODONTIDAE

†Family ICHTHYODECTIDAE





†Order TSELFATIIFORMES †Family Plethodidae (= Bananogmiidae)

†Family Protobramidae

†Family Tselfatiidae





†Order ARARIPICHTHYIFORMES †Family ARARIPICHTHYIDAE





Supercohort TELEOCEPHALA—crown-group Teleostei

Cohort ELOPOMORPHA Order ELOPIFORMES (23)—tenpounders Family ELOPIDAE (68)—tenpounders (ladyfishes)

Family MEGALOPIDAE (69)—tarpons





Order ALBULIFORMES (24)—bonefishes Family ALBULIDAE (70)—bonefishes





Order NOTACANTHIFORMES (25)—halosaurs and deep-sea spiny eels Family HALOSAURIDAE (71)—halosaurs

Family NOTACANTHIDAE (72)—deep-sea spiny eels





Order ANGUILLIFORMES (Apodes) (26)—eels Family PROTANGUILLIDAE (73)—primitive cave eels

Family SYNAPHOBRANCHIDAE (74)—cutthroat eels

Family HETERENCHELYIDAE (75)—mud eels

Family MYROCONGRIDAE (76)—myroconger eels

Family MURAENIDAE (Heteromyridae) (77)—moray eels

Family CHLOPSIDAE (Xenocongridae) (78)—false morays

Family DERICHTHYIDAE (79) (includes Colocongridae)—longneck or narrowneck and shorttail eels

Family OPHICHTHIDAE (80)—snake eels and worm eels

Family MURAENESOCIDAE (81)—pike congers

Family NETTASTOMATIDAE (82)—duckbill eels

Family CONGRIDAE (83)—conger eels

Family MORINGUIDAE (84)—spaghetti eels

Family CYEMATIDAE (85)—bobtail snipe eels

Family MONOGNATHIDAE (86)—onejaw gulpers

Family SACCOPHARYNGIDAE (87)—swallowers

Family EURYPHARYNGIDAE (88)—gulpers or pelican eels

Family NEMICHTHYIDAE (89)—snipe eels

Family SERRIVOMERIDAE (90)—sawtooth eels

Family ANGUILLIDAE (91)—freshwater eels





OSTEOGLOSSOCEPHALA

Cohort OSTEOGLOSSOMORPHA †Order LYCOPTERIFORMES †Family LYCOPTERIDAE





Order HIODONTIFORMES (27)—mooneyes Family HIODONTIDAE (92)—mooneyes





Order OSTEOGLOSSIFORMES (28)—bonytongues Family PANTODONTIDAE (93)—butterflyfishes

Family OSTEOGLOSSIDAE (94)—osteoglossids or bonytongues

Family NOTOPTERIDAE (95)

Family MORMYRIDAE (96)—elephantfishes

Family GYMNARCHIDAE (97)—abas





CLUPEOCEPHALA

Cohort OTOCEPHALA (= OSTARIOCLUPEOMORPHA, OTOMORPHA)

Superorder CLUPEOMORPHA †Order ELLIMMICHTHYIFORMES †Family ARMIGATIDAE

†Family SORBINICHTHYIDAE

†Family PARACLUPEIDAE (= Ellimmichthyidae)





Order CLUPEIFORMES (29)—herrings Family DENTICIPITIDAE (98)—denticle herrings

Family PRISTIGASTERIDAE (99)—longfin herrings

Family ENGRAULIDAE (Engraulididae and Stolephoridae) (100)—anchovies

Family CHIROCENTRIDAE (101)—wolf herrings

Family CLUPEIDAE (102)—herrings (shads, sprats, sardines, pilchards, and menhadens)





Superorder ALEPOCEPHALI Order ALEPOCEPHALIFORMES (30)—slickheads and tubeshoulders Family PLATYTROCTIDAE (Searsiidae) (103)—tubeshoulders

Family BATHYLACONIDAE (104)—bathylaconids

Family ALEPOCEPHALIDAE (105)—slickheads





Superorder OSTARIOPHYSI Series ANOTOPHYSI

Order GONORYNCHIFORMES (31)—milkfishes Family CHANIDAE (106)—milkfishes

Family GONORYNCHIDAE (107)—beaked sandfishes

Family KNERIIDAE (108)—knerias and snake mudheads





Series OTOPHYSI

Subseries Cypriniphysi

Order CYPRINIFORMES (32)—carps, loaches, minnows, and relatives Family CYPRINIDAE (109)—minnows, carps, and loaches

Family PSILORHYNCHIDAE (110)—mountain carps

†Family JIANGHANICHTHYIDAE

Family GYRINOCHEILIDAE (111)—algae eaters

Family CATOSTOMIDAE (112)—suckers

Family BOTIIDAE (113)—botiid loaches

Family VAILLANTELLIDAE (114)—long-fin loaches

Family COBITIDAE (115)—loaches

Family BALITORIDAE (Homalopteridae) (116)—hillstream or river loaches

Family GASTROMYZONTIDAE (117)—gastromyzontid or sucker loaches

Family NEMACHEILIDAE (118)—stone loaches

Family BARBUCCIDAE (119)—fire-eyed loaches

Family ELLOPOSTOMATIDAE (120)—sturgeon-mouthed loaches

Family SERPENTICOBITIDAE (121)—serpent loaches





Subseries Characiphysi

Order CHARACIFORMES (33)—characins Family DISTICHODONTIDAE (122)—distichodontids

Family CITHARINIDAE (123)—citharinids

Family CRENUCHIDAE (124)—South American darters

Family ALESTIDAE (ALESTIIDAE) (125)—African tetras

Family HEPSETIDAE (126)—African pikes

Family ERYTHRINIDAE (127)—trahiras

Family PARODONTIDAE (128)—parodontids

Family CYNODONTIDAE (129)—cynodontids

Family SERRASALMIDAE (130)—pacus, silver dollars, and piranhas

Family HEMIODONTIDAE (131)—hemiodontids

Family ANOSTOMIDAE (132)—toothed headstanders

Family CHILODONTIDAE (133)—headstanders

Family CURIMATIDAE (134)—toothless characiforms

Family PROCHILODONTIDAE (135)—flannel-mouth characiforms

Family LEBIASINIDAE (136)—pencil fishes

Family CTENOLUCIIDAE (137)—pike-characids

Family ACESTRORHYNCHIDAE (138)—acestrorhynchids

Family CHARACIDAE (139)—characins

Family GASTEROPELECIDAE (140)—freshwater hatchetfishes





Subseries Siluriphysi

Order SILURIFORMES (Nematognathi) (34)—catfishes †Family ANDINICHTHYIDAE

†Family BACHMANNIIDAE

Family DIPLOMYSTIDAE (141)—velvet catfishes

†Family HYPSIDORIDAE

Family CETOPSIDAE (142)—whale catfishes

Family TRICHOMYCTERIDAE (Pygidiidae) (143)—pencil catfishes or parasitic catfishes

Family NEMATOGENYIDAE (144)—mountain catfishes

Family CALLICHTHYIDAE (145)—callichthyid armored catfishes

Family SCOLOPLACIDAE (146)—spiny dwarf catfishes

Family ASTROBLEPIDAE (Argidae) (147)—climbing catfishes

Family LORICARIIDAE (148)—suckermouth armored catfishes

Family SILURIDAE (149)—sheatfishes

Family AUSTROGLANIDIDAE (150)—rock catfishes

Family PANGASIIDAE (151)—shark catfishes

Family CHACIDAE (152)—squarehead, angler, or frogmouth catfishes

Family PLOTOSIDAE (153)—eeltail catfishes

Family RITIDAE (154)—ritas and nanobagrids

Family AILIIDAE (155)—Asian “schilbeids”

Family HORABAGRIDAE (156)—imperial or sun catfishes

Family BAGRIDAE (157)—bagrid catfishes

Family AKYSIDAE (158)—stream catfishes

Family AMBLYCIPITIDAE (159)—torrent catfishes

Family SISORIDAE (Bagariidae) (160)—sisorid catfishes

Family ERETHISTIDAE (161)—erethistid catfishes

Family AMPHILIIDAE (162)—loach catfishes

Family MALAPTERURIDAE (163)—electric catfishes

Family MOCHOKIDAE (164)—squeakers or upside-down catfishes

Family SCHILBEIDAE (SCHILBIDAE) (165)—schilbeid or butter catfishes

Family AUCHENOGLANIDIDAE (166)—auchenoglanidids

Family CLAROTEIDAE (167)—claroteids

Family LACANTUNIIDAE (168)—Chiapas catfishes

Family CLARIIDAE (169)—airbreathing catfishes

Family HETEROPNEUSTIDAE (Saccobranchidae) (170)—airsac catfishes

Family ANCHARIIDAE (171)—Malagasy catfishes

Family ARIIDAE (Tachysuridae) (172)—sea catfishes

Family ASPREDINIDAE (173)—banjo catfishes

Family DORADIDAE (174)—thorny catfishes

Family AUCHENIPTERIDAE (175)—driftwood catfishes

Family CRANOGLANIDIDAE (176)—armorhead catfishes

Family ICTALURIDAE (Ameiuridae) (177)—North American catfishes

Family HEPTAPTERIDAE (178)—heptapterids

Family PIMELODIDAE (179)—long-whiskered catfishes

Family PSEUDOPIMELODIDAE (180)—bumblebee catfishes





Order GYMNOTIFORMES (35)—Neotropical knifefishes Family GYMNOTIDAE (181)—nakedback knifefishes

Family RHAMPHICHTHYIDAE (182)—sand knifefishes

Family HYPOPOMIDAE (183)—bluntnose knifefishes

Family STERNOPYGIDAE (184)—glass knifefishes

Family APTERONOTIDAE (185)—ghost knifefishes





Cohort EUTELEOSTEI Order LEPIDOGALAXIIFORMES (36)—salamanderfishes Family LEPIDOGALAXIIDAE (186)—salamanderfishes





Superorder PROTACANTHOPTERYGII Order SALMONIFORMES (37)—trout, salmon, and whitefish Family SALMONIDAE (187)—trout, salmon, and whitefish





Order ESOCIFORMES (Haplomi, Esocae) (38)—pikes and mudminnows †Family PALAEOESOCIDAE

Family ESOCIDAE (188)—pikes

Family UMBRIDAE (189)—mudminnows





Superorder OSMEROMORPHA Order ARGENTINIFORMES (39)—marine smelts Family ARGENTINIDAE (190)—argentines or herring smelts

Family OPISTHOPROCTIDAE (191)—barreleyes or spookfishes

Family MICROSTOMATIDAE (192)—pencilsmelts

Family BATHYLAGIDAE (193)—deepsea smelts





Order GALAXIIFORMES (40)—galaxiiforms Family GALAXIIDAE (194)—galaxiids





Order OSMERIFORMES (41)—freshwater smelts Family OSMERIDAE (195)—Northern Hemisphere smelts

Family PLECOGLOSSIDAE (196)—Ayu or sweetfish

Family SALANGIDAE (197)—icefishes or noodlefishes

Family PROTOTROCTIDAE (198)—southern graylings

Family RETROPINNIDAE (199)—southern smelts





Order STOMIIFORMES (Stomiatiformes) (42)—dragonfishes Family GONOSTOMATIDAE (200)—bristlemouths

Family STERNOPTYCHIDAE (201)—marine hatchetfishes

Family PHOSICHTHYIDAE (Photichthyidae) (202)—lightfishes

Family STOMIIDAE (203)—barbeled dragonfishes





Superorder ATELEOPODOMORPHA Order ATELEOPODIFORMES (43)—jellynose fishes Family ATELEOPODIDAE (204)—jellynose fishes





Superorder CYCLOSQUAMATA Order AULOPIFORMES (44)—lizardfishes †Family ICHTHYOTRINGIDAE

†Family DERCETIDAE

†Family PRIONOLEPIDIDAE

†Family HALECIDAE

Family SYNODONTIDAE (205)—lizardfishes

Family AULOPIDAE (206)—flagfins

Family PSEUDOTRICHONOTIDAE (207)—sand-diving lizardfishes

Family PARAULOPIDAE (208)—cucumber fishes

Family IPNOPIDAE (209)—deepsea tripod fishes

Family GIGANTURIDAE (210)—telescopefishes

Family BATHYSAUROIDIDAE (211)—largescale deep-sea lizardfish

Family BATHYSAURIDAE (212)—deepsea lizardfishes

Family CHLOROPHTHALMIDAE (213)—greeneyes

Family NOTOSUDIDAE (Scopelosauridae) (214)—waryfishes

Family SCOPELARCHIDAE (215)—pearleyes

Family EVERMANNELLIDAE (216)—sabertooth fishes

Family SUDIDAE (217)—sudid barracudinas

Family PARALEPIDIDAE (218)—barracudinas

†Family Enchodontidae

Family ALEPISAURIDAE (219)—lancetfishes

Family LESTIDIIDAE (220)—naked barracudinas





Superorder SCOPELOMORPHA Order MYCTOPHIFORMES (45)—lanternfishes Family NEOSCOPELIDAE (221)—blackchins

Family MYCTOPHIDAE (222)—lanternfishes





†Order CTENOTHRISSIFORMES





Superorder LAMPRIMORPHA Order LAMPRIFORMES (Lampridiformes, Allotriognathi) (46)—opahs Family VELIFERIDAE (223)—velifers

Family LAMPRIDAE (Lamprididae) (224)—opahs

Family LOPHOTIDAE (225)—crestfishes

Family RADIICEPHALIDAE (226)—tapertails

Family TRACHIPTERIDAE (227)—ribbonfishes

Family REGALECIDAE (228)—oarfishes





Superorder PARACANTHOPTERYGII Order POLYMIXIIFORMES (47)—beardfishes Family POLYMIXIIDAE (229)—beardfishes





†Order SPHENOCEPHALIFORMES †Family SPHENOCEPHALIDAE





Order PERCOPSIFORMES (48)—trout-perches Family PERCOPSIDAE (230)—trout-perches

†Family MCCONICHTHYIDAE

Family APHREDODERIDAE (231)—pirate perches

Family AMBLYOPSIDAE (232)—cavefishes





Order ZEIFORMES (49)—dories Family CYTTIDAE (233)—lookdown dories

Family OREOSOMATIDAE (234)—oreos

Family PARAZENIDAE (235)—smooth dories

Family ZENIONTIDAE (Zenionidae) (236)—armoreye dories

Family GRAMMICOLEPIDIDAE (237)—tinselfishes

Family ZEIDAE (238)—dories





Order STYLEPHORIFORMES (50)—tube-eyes or thread-tails Family STYLEPHORIDAE (239)—tube-eyes or thread-tails





Order GADIFORMES (51)—cods and hakes Family MELANONIDAE (240)—pelagic cods

Family STEINDACHNERIIDAE (241)—luminous and southern hakes

Family BATHYGADIDAE (242)—rattails

Family MACROURIDAE (Coryphaenoididae) (243)—grenadiers (rattails)

Family TRACHYRINCIDAE (244)—whiptails and trachyrincines

Family EUCLICHTHYIDAE (245)—eucla cod

Family MORIDAE (246)—deepsea cods, codlings, hakelings

Family MACRURONIDAE (247)—southern hakes

Family MERLUCCIIDAE (248)—merlucciid hakes

Family RANICIPITIDAE (249)—tadpole cods

Family BREGMACEROTIDAE (250)—codlets

Family MURAENOLEPIDIDAE (251)—eel cods and moray cods

Family GADIDAE (252)—cods





Superorder ACANTHOPTERYGII Series BERYCIDA

Order HOLOCENTRIFORMES (52)—squirrelfishes Family HOLOCENTRIDAE (253)—squirrelfishes





Order TRACHICHTHYIFORMES (53)—roughies Family ANOPLOGASTRIDAE (254)—fangtooths

Family DIRETMIDAE (255)—spinyfins

Family ANOMALOPIDAE (256)—flashlight or lanterneye fishes

†Family PSEUDOMONOCENTRIDAE (Pseudomonocentrididae)

Family MONOCENTRIDAE (Monocentrididae) (257)—pinecone fishes

Family TRACHICHTHYIDAE (258)—roughies





Order BERYCIFORMES (54)—beryciforms Family GIBBERICHTHYIDAE (259)—gibberfishes

Family STEPHANOBERYCIDAE (260)—pricklefishes

Family HISPIDOBERYCIDAE (261)—hispidoberycids

Family RONDELETIIDAE (262)—redmouth whalefishes

Family BARBOURISIIDAE (263)—red (redvelvet) whalefishes

Family CETOMIMIDAE (264)—flabby whalefishes

Family MELAMPHAIDAE (265)—bigscale fishes

Family BERYCIDAE (266)—alfonsinos





Series PERCOMORPHA

Subseries Ophidiida

Order OPHIDIIFORMES (55)—cusk-eels Family CARAPIDAE (267)—pearlfishes

Family OPHIDIIDAE (268)—cusk-eels

Family BYTHITIDAE (269)—viviparous brotulas

Family APHYONIDAE (270)—aphyonids, blind cusk-eels

Family PARABROTULIDAE (271)—false brotulas





Subseries Batrachoidida

Order BATRACHOIDIFORMES (Haplodoci) (56)—toadfishes Family BATRACHOIDIDAE (272)—toadfishes





Subseries Gobiida

Order KURTIFORMES (57)—nurseryfishes and cardinalfishes Family KURTIDAE (273)—nurseryfishes

Family APOGONIDAE (274)—cardinalfishes





Order GOBIIFORMES (58)—gobies Family RHYACICHTHYIDAE (275)—loach gobies

Family ODONTOBUTIDAE (276)—freshwater sleepers

Family MILYERINGIDAE (277)—blind cave gobies

Family ELEOTRIDAE (Eleotrididae) (278)—spinycheek sleepers

Family BUTIDAE (279)—butid sleepers

Family THALASSELEOTRIDIDAE (280)—ocean sleepers

Family OXUDERCIDAE (= GOBIONELLIDAE) (281)—Gobionellus-like and mudskipper gobies

Family GOBIIDAE (282)—gobies





Subseries Ovalentaria Family AMBASSIDAE (Chandidae) (283)—Asiatic glassfishes

Family EMBIOTOCIDAE (284)—surfperches

Family GRAMMATIDAE (Grammidae) (285)—basslets

Family PLESIOPIDAE (286)—roundheads

Family POLYCENTRIDAE (287)—South American leaffishes

Family POMACENTRIDAE (288)—damselfishes

Family PSEUDOCHROMIDAE (289)—dottybacks

Family OPISTOGNATHIDAE (290)—jawfishes





Order MUGILIFORMES (59)—mullets Family MUGILIDAE (291)—mullets





Order CICHLIFORMES (60)—cichlids and convict blennies Family CICHLIDAE (292)—cichlids

Family PHOLIDICHTHYIDAE (293)—convict blenny





Order BLENNIIFORMES (61)—Blennies Family TRIPTERYGIIDAE (294)—triplefin blennies

Family DACTYLOSCOPIDAE (295)—sand stargazers

Family BLENNIIDAE (296)—combtooth blennies

Family CLINIDAE (297)—kelp blennies

Family LABRISOMIDAE (298)—labrisomid blennies

Family CHAENOPSIDAE (299)—tube blennies





Order GOBIESOCIFORMES (62)—clingfishes Family GOBIESOCIDAE (300)—clingfishes





Order ATHERINIFORMES (63)—silversides Family ATHERINOPSIDAE (301)—New World silversides

Family NOTOCHEIRIDAE (302)—surf silversides

Family ISONIDAE (303)—surf sardines

Family MELANOTAENIIDAE (304)—rainbowfishes and blue eyes

Family ATHERIONIDAE (305)—pricklenose silversides

Family DENTATHERINIDAE (306)—Mercer's tusked silverside

Family PHALLOSTETHIDAE (307)—priapiumfishes

Family ATHERINIDAE (308)—Old World silversides





Order BELONIFORMES (64)—needlefishes Family ADRIANICHTHYIDAE (309)—adrianichthyids

Family EXOCOETIDAE (310)—flyingfishes

Family HEMIRAMPHIDAE (311)—halfbeaks

Family ZENARCHOPTERIDAE (312)—viviparous halfbeaks

Family BELONIDAE (313)—needlefishes

Family SCOMBERESOCIDAE (314)—sauries





Order CYPRINODONTIFORMES (Microcyprini) (65)—killifishes Family APLOCHEILIDAE (315)—Asian rivulines

Family NOTHOBRANCHIIDAE (316)—African rivulines

Family RIVULIDAE (317)—New World rivulines

Family PROFUNDULIDAE (318)—Middle American killifishes

Family GOODEIDAE (319)—goodeids

Family FUNDULIDAE (320)—topminnows

Family VALENCIIDAE (321)—Valencia toothcarps

Family CYPRINODONTIDAE (322)—pupfishes

Family ANABLEPIDAE (323)—four-eyed fishes

Family POECILIIDAE (324)—livebearers





Order SYNBRANCHIFORMES (66)—swamp eels Family SYNBRANCHIDAE (325)—swamp eels

Family CHAUDHURIIDAE (326)—earthworm eels

Family MASTACEMBELIDAE (327)—freshwater spiny eels





Order CARANGIFORMES (67)—jacks Family NEMATISTIIDAE (328)—roosterfishes

Family CORYPHAENIDAE (329)—dolphinfishes

Family RACHYCENTRIDAE (330)—cobias

Family ECHENEIDAE (Echeneididae) (331)—remoras (sharksuckers)

Family CARANGIDAE (332)—jacks and pompanos

Family MENIDAE (333)—moonfishes





Order ISTIOPHORIFORMES (68)—barracudas and billfishes Family SPHYRAENIDAE (334)—barracudas

Family XIPHIIDAE (335)—swordfishes

Family ISTIOPHORIDAE (336)—billfishes





Order ANABANTIFORMES (Labyrinthici) (69)—labyrinth fishes Family ANABANTIDAE (337)—climbing gouramies

Family HELOSTOMATIDAE (338)—kissing gouramies

Family OSPHRONEMIDAE (339)—gouramies and fighting fishes

Family CHANNIDAE (340)—snakeheads

Family NANDIDAE (341)—Asian leaffishes

Family BADIDAE (342)—chameleonfishes

Family PRISTOLEPIDIDAE (343)—(Malayan) leaffishes





Order PLEURONECTIFORMES (Heterosomata) (70)—flatfishes Family PSETTODIDAE (344)—spiny turbots

Family CITHARIDAE (345)—largescale flounders

Family SCOPHTHALMIDAE (346)—turbots

Family PARALICHTHYIDAE (347)—sand flounders

Family PLEURONECTIDAE (348)—righteye flounders

Family BOTHIDAE (349)—lefteye flounders

Family PARALICHTHODIDAE (350)—measles or peppered flounders

Family POECILOPSETTIDAE (351)—bigeye flounders

Family RHOMBOSOLEIDAE (352)—rhombosoleids

Family ACHIROPSETTIDAE (353)—southern flounders

Family SAMARIDAE (354)—crested flounders

Family ACHIRIDAE (355)—American soles

Family SOLEIDAE (356)—soles

Family CYNOGLOSSIDAE (357)—tonguefishes





Order SYNGNATHIFORMES (71)—pipefishes and seahorses Family PEGASIDAE (358)—seamoths

Family SOLENOSTOMIDAE (359)—ghost pipefishes

Family SYNGNATHIDAE (360)—pipefishes and seahorses

Family AULOSTOMIDAE (361)—trumpetfishes

Family FISTULARIIDAE (362)—cornetfishes

Family MACRORAMPHOSIDAE (363)—snipefishes

Family CENTRISCIDAE (364)—shrimpfishes

Family DACTYLOPTERIDAE (Cephalacanthidae) (365)—flying gurnards





Order ICOSTEIFORMES (Malacichthyes) (72)—ragfishes Family ICOSTEIDAE (366)—ragfishes





Order CALLIONYMIFORMES (73) Family CALLIONYMIDAE (367)—dragonets

Family DRACONETTIDAE (368)—slope dragonets





Order SCOMBROLABRACIFORMES (74)—longfin escolars Family SCOMBROLABRACIDAE (369)—longfin escolars





Order SCOMBRIFORMES (Pelagia) (75)—mackerels Family GEMPYLIDAE (370)—snake mackerels

Family TRICHIURIDAE (371)—cutlassfishes

Family SCOMBRIDAE (372)—mackerels and tunas

Family AMARSIPIDAE (373)—amarsipas

Family CENTROLOPHIDAE (374)—medusafishes

Family NOMEIDAE (375)—driftfishes

Family ARIOMMATIDAE (376)—ariommatids

Family TETRAGONURIDAE (377)—squaretails

Family STROMATEIDAE (378)—butterfishes





Order TRACHINIFORMES (Uranoscopiformes in part) (76) Family CHIASMODONTIDAE (379)—swallowers

Family CHAMPSODONTIDAE (380)—gapers

Family PINGUIPEDIDAE (381)—sandperches

Family CHEIMARRHICHTHYIDAE (382)—New Zealand torrentfishes

Family TRICHONOTIDAE (383)—sanddivers

Family CREEDIIDAE (384)—sandburrowers

Family PERCOPHIDAE (385)—duckbills

Family LEPTOSCOPIDAE (386)—southern sandfishes

Family AMMODYTIDAE (387)—sand lances

Family TRACHINIDAE (388)—weeverfishes

Family URANOSCOPIDAE (389)—stargazers





Order LABRIFORMES (77)—wrasses and relatives Family LABRIDAE (390)—wrasses

Family ODACIDAE (391)—cales

Family SCARIDAE (Callyodontidae) (392)—parrotfishes





Order PERCIFORMES (78)—perches Family CENTROPOMIDAE (393)—snooks

Family LATIDAE (394)—lates perches

Family GERREIDAE (395)—mojarras

Family CENTROGENYIDAE (396)—false scorpionfishes

Family PERCILIIDAE (397)—southern basses

Family HOWELLIDAE (398)—oceanic basslets

Family ACROPOMATIDAE (399)—lanternbellies

Family EPIGONIDAE (400)—deepwater cardinalfishes

Family POLYPRIONIDAE (401)—wreckfishes

Family LATEOLABRACIDAE (402)—Asian seaperches

Family MULLIDAE (403)—goatfishes

Family GLAUCOSOMATIDAE (404)—pearl perches

Family PEMPHERIDAE (405)—sweepers

Family OPLEGNATHIDAE (406)—knifejaws

Family KUHLIIDAE (407)—flagtails

Family BATHYCLUPEIDAE (408)—bathyclupeids

Family TOXOTIDAE (409)—archerfishes

Family ARRIPIDAE (410)—Australasian salmon (kahawai)

Family DICHISTIIDAE (Coracinidae) (411)—galjoen fishes

Family KYPHOSIDAE (412)—sea chubs

Family TERAPONTIDAE (Teraponidae, Theraponidae) (413)—grunters or tigerperches

Family PERCICHTHYIDAE (433)—temperate perches

Family SINIPERCIDAE (415)—Chinese perches

Family ENOPLOSIDAE (416)—oldwives

Family PENTACEROTIDAE (417)—armorheads

Family DINOPERCIDAE (418)—cavebasses

Family BANJOSIDAE (419)—banjofishes

Family CENTRARCHIDAE (420)—sunfishes

Family SERRANIDAE (421)—sea basses

Family PERCIDAE (422)—perches

Family LACTARIIDAE (423)—false trevallies

Family DINOLESTIDAE (424)—long-finned pikes

Family SCOMBROPIDAE (425)—gnomefishes

Family POMATOMIDAE (426)—bluefishes

Family BRAMIDAE (427)—pomfrets

Family CARISTIIDAE (428)—manefishes

Family MONODACTYLIDAE (429)—moonfishes (fingerfishes)

Family PRIACANTHIDAE (430)—bigeyes (catalufas)

Family LEIOGNATHIDAE (431)—ponyfishes, slimys, or slipmouths

Family CHAETODONTIDAE (432)—butterflyfishes

Family POMACANTHIDAE (433)—angelfishes

Family MALACANTHIDAE (434)—tilefishes

Family HAEMULIDAE (Pomadasyidae) (435)—grunts

Family HAPALOGENIIDAE (436)—barbeled grunters

Family LUTJANIDAE (437)—snappers

Family CAESIONIDAE (438)—fusiliers

Family CIRRHITIDAE (439)—hawkfishes

Family CHIRONEMIDAE (440)—kelpfishes

Family APLODACTYLIDAE (Haplodactylidae) (441)—marblefishes

Family CHEILODACTYLIDAE (442)—morwongs

Family LATRIDAE (443)—trumpeters

Family CEPOLIDAE (444)—bandfishes

Family SCATOPHAGIDAE (445)—scats

Family SIGANIDAE (446)—rabbitfishes

Family BOVICHTIDAE (Bovichthyidae) (447)—temperate icefishes

Family PSEUDAPHRITIDAE (448)—catadromous icefishes

Family ELEGINOPSIDAE (449)—Patagonian blennies

Family NOTOTHENIIDAE (450)—cod icefishes

Family HARPAGIFERIDAE (451)—spiny plunderfishes

Family ARTEDIDRACONIDAE (452)—barbeled plunderfishes

Family BATHYDRACONIDAE (453)—Antarctic dragonfishes

Family CHANNICHTHYIDAE (Chaenichthyidae) (454)—crocodile icefishes





Order SCORPAENIFORMES (79)—mail-cheeked fishes Family SCORPAENIDAE (455)—scorpionfishes (rockfishes)

Family APLOACTINIDAE (456)—velvetfishes

Family ESCHMEYERIDAE (457)—cofishes

Family PATAECIDAE (458)—Australian prowfishes

Family GNATHANACANTHIDAE (459)—red velvetfishes

Family CONGIOPODIDAE (460)—racehorses (pigfishes or horsefishes)

Family TRIGLIDAE (461)—searobins (gurnards)

Family PERISTEDIIDAE (462)—armored searobins

Family BEMBRIDAE (463)—deepwater flatheads

Family PLATYCEPHALIDAE (464)—flatheads

Family HOPLICHTHYIDAE (465)—ghost flatheads

Family NORMANICHTHYIDAE (466)—barehead scorpionfishes or mote sculpins

Family BATHYMASTERIDAE (467)—ronquils

Family EULOPHIIDAE (468)—eulophiids

Family ZOARCIDAE (469)—eelpouts

Family STICHAEIDAE (470)—pricklebacks

Family CRYPTACANTHODIDAE (471)—wrymouths

Family PHOLIDAE (472)—gunnels

Family ANARHICHADIDAE (473)—wolffishes

Family PTILICHTHYIDAE (474)—quillfishes

Family ZAPRORIDAE (475)—prowfishes

Family SCYTALINIDAE (476)—graveldivers

Family HYPOPTYCHIDAE (477)—sand eel

Family AULORHYNCHIDAE (478)—tubesnouts

Family GASTEROSTEIDAE (479)—sticklebacks

Family INDOSTOMIDAE (480)—armored sticklebacks

Family ANOPLOPOMATIDAE (481)—sablefishes

Family ZANIOLEPIDIDAE (482)—combfishes

Family HEXAGRAMMIDAE (483)—greenlings

Family TRICHODONTIDAE (484)—sandfishes

Family JORDANIIDAE (485)—longfin sculpin

Family RHAMPHOCOTTIDAE (EREUNIIDAE) (486)—grunt sculpins

Family SCORPAENICHTHYIDAE (487)—Cabezon

Family AGONIDAE (HEMITRIPTERIDAE) (488)—poachers and searavens

Family COTTIDAE (489)—sculpins

Family PSYCHROLUTIDAE (490)—fathead sculpins

Family BATHYLUTICHTHYIDAE (491)—Antarctic sculpins

Family CYCLOPTERIDAE (492)—lumpfishes (lumpsuckers)

Family LIPARIDAE (Liparididae) (493)—snailfishes





Order MORONIFORMES (80)—temperate basses Family MORONIDAE (494)—temperate basses

Family DREPANEIDAE (495)—sicklefishes

Family EPHIPPIDAE (496)—spadefishes





Order ACANTHURIFORMES—(81) surgeonfishes and relatives Family EMMELICHTHYIDAE (497)—rovers

Family SCIAENIDAE (498)—drums (croakers)

Family LUVARIDAE (499)—louvar

Family ZANCLIDAE (500)—Moorish Idols

†Family MASSALONGIIDAE

Family ACANTHURIDAE (501)—surgeonfishes





Order SPARIFORMES (82)—breams and porgies Family CALLANTHIIDAE (502)—splendid perches or groppos

Family SILLAGINIDAE (503)—sillagos (whitings, smelt-whitings)

Family LOBOTIDAE (504)—tripletails

Family NEMIPTERIDAE (505)—threadfin breams

Family LETHRINIDAE (506)—emperors or emperor breams

Family SPARIDAE (507)—porgies





Order CAPROIFORMES (83)—boarfishes Family CAPROIDAE (508)—boarfishes





Order LOPHIIFORMES (84)—anglerfishes Family LOPHIIDAE (509)—goosefishes

Family ANTENNARIIDAE (510)—frogfishes

Family TETRABRACHIIDAE (511)—tetrabrachiid frogfishes

Family LOPHICHTHYIDAE (512)—lophichthyid frogfishes

Family BRACHIONICHTHYIDAE (513)—handfishes (warty anglers)

Family CHAUNACIDAE (514)—coffinfishes or sea toads

Family OGCOCEPHALIDAE (515)—batfishes

Family CAULOPHRYNIDAE (516)—fanfins

Family NEOCERATIIDAE (517)—spiny seadevils

Family MELANOCETIDAE (518)—black seadevils

Family HIMANTOLOPHIDAE (519)—footballfishes

Family DICERATIIDAE (520)—double anglers

Family ONEIRODIDAE (521)—dreamers

Family THAUMATICHTHYIDAE (522)—wolftrap anglers

Family CENTROPHRYNIDAE (523)—prickly seadevils

Family CERATIIDAE (524)—warty seadevils

Family GIGANTACTINIDAE (525)—whipnose anglers

Family LINOPHRYNIDAE (526)—leftvents





Order TETRAODONTIFORMES (Plectognathi) (85)—plectognaths Family TRIODONTIDAE (527)—threetooth puffers

Family TRIACANTHODIDAE (528)—spikefishes

Family TRIACANTHIDAE (529)—triplespines

Family ARACANIDAE (530)—deepwater boxfishes

Family OSTRACIIDAE (Ostraciontidae) (531)—boxfishes (cowfishes and trunkfishes)

Family BALISTIDAE (532)—triggerfishes

Family MONACANTHIDAE (533)—filefishes

Family MOLIDAE (534)—molas (ocean sunfishes)

Family TETRAODONTIDAE (535)—puffers

Family DIODONTIDAE (536)—porcupinefishes (burrfishes)





Bibliography

Index

End User License Agreement





Fishes of the World


Fifth Edition




Joseph S. Nelson

Terry C. Grande

Mark V. H. Wilson





Cover image: Mark V. H. Wilson

Cover design: Wiley

Copyright © 2016 by John Wiley & Sons, Inc. All rights reserved.

Published by John Wiley & Sons, Inc., Hoboken, New Jersey.

Published simultaneously in Canada.

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Library of Congress Cataloging-in-Publication Data:

Names: Nelson, Joseph S., author. | Grande, Terry, author. | Wilson, Mark V. H., author.

Title: Fishes of the world / Joseph S. Nelson, Terry C. Grande, Mark V. H. Wilson.

Description: Fifth edition. | Hoboken, New Jersey : John Wiley & Sons, 2016. | Includes bibliographical references and index.

Identifiers: LCCN 2015037522 | ISBN 9781118342336 (cloth); 9781119220824 (ebk.); 9781119220817 (ebk.); 9781119174844 (ebk.)

Subjects: LCSH: Fishes—Classification.

Classification: LCC QL618 .N4 2016 | DDC 597.01/2–dc23 LC record available at http://lccn.loc.gov/2015037522





Memories of Joe Nelson


Terry Grande and Mark Wilson asked if I could write a short essay involving my own personal memories of Joe, and I willingly agreed to do so. It should be noted that the paragraphs below are not an in-depth summary of Joe's life, but rather personal recollections. A more complete summary of his most interesting life and highly productive scientific career, written by David G. Smith for the continuing series entitled “Historical Perspectives,” appeared recently on pages 169–176 of Copeia, 2011, no. 4.



In 1976, Joseph S. Nelson published the first edition of his book, Fishes of the World, involving all living and extinct groups of fishes. The work was widely recognized as the definitive source of information on fish classification, and a “must have” reference in the library of any person seriously involved in work with this group of animals. The success of this book, coupled with ongoing changes in knowledge and understanding of fish classification, brought the realization that periodic updates to this book would be necessary. This resulted in new editions during each succeeding decade (1984, 1994, and 2006), and it was Joe's intention that these would be followed by another edition during the succeeding decade. However, Joe's illness and subsequent passing made it impossible for him to finish the project alone, and led him to invite Terry Grande and Mark Wilson to assist him, and then, when his illness became terminal, to complete the project without him.

I first met Joe Nelson, during the 1970s, at one of the annual meetings of our primary professional society, the American Society of Ichthyologists and Herpetologists (ASIH). I was already aware of his work on salmonid fishes, nine-spine sticklebacks, and New Zealand marine fishes through the exchange of reprints of publications, and particularly his landmark 1976 work, the first edition of Fishes of the World, which had been accepted by ichthyologists (and scientists in general) as the most important published modern summary on classification of the world's living and fossil fishes. This publication had already propelled Joe, at the relatively young age of 39, to the forefront among world ichthyologists.

Although I knew Joe only casually in those early days, we nevertheless had frequent opportunities to discuss matters of mutual interest, especially those involving taxonomic and distributional problems involving North American marine and freshwater fishes. He immediately struck me as a friendly and soft-spoken person, and certainly not one to make a big deal of his already impressive professional accomplishments. This impression was one I never had any reason to change and, it can be said without fear of contradiction, one that has always been shared by all his colleagues and friends.

An incident occurred at a social gathering at one of these meetings when a young student, upon seeing the name on his name tag, approached Joe and rather tentatively asked, “Are you the Joe Nelson?” His positive response elicited an expression of awe, and in subsequent years this incident led to our referring to Joe as “The Joe Nelson.”

As noted, Joe and I were regular attendees at the annual ASIH meetings. We were sometimes accompanied by our respective spouses, Claudine and Nancy, both of whom were more interested in the attractions and activities associated with the local meeting site than sitting in a room listening to presentations on fishes. The 1988 ASIH meeting in Ann Arbor, Michigan, was one such occasion, and that meeting can be pinpointed as the beginning of a long and enduring friendship. In subsequent years, when the question arose whether Nancy might choose to accompany me to the meetings, the deciding factor was whether Joe and Claudine would be there. This friendship resulted in trips together to rather exotic meeting sites such as La Paz, Baja California (Mexico), in 2000, and Manaus, Brazil, in 2003.

In 1991, Joe was asked to serve as chair of the joint AFS (American Fisheries Society)–ASIH committee on common and scientific names of fishes (often abbreviated to “c & sn of fishes”). This committee, which had been in existence since the late 1940s, is charged with maintaining and publishing, at approximate ten-year intervals, an updated checklist, with broad indications of distributions, of all species of United States and Canadian freshwater fishes and of marine species occurring within the 200-meter depth limit. The geographical scope of the list was expanded in the sixth edition, in 2004, to include Mexico as well. This list serves as an important authoritative reference for taxonomists, fisheries biologists, aquarists, and those involved in matters of conservation. Equally important are the accompanying appendices, which include explanations and justifications for changes or additions to the list. Throughout the preceding years, I had regularly provided pertinent information to the committee as it came to my attention. Shortly after assuming the chair, Joe invited me to join the committee on a formal basis.

Joe's years of productive ichthyological research had resulted, by the turn of the century, in well over one hundred papers in professional journals, together with books such as the definitive regional work co-authored with Martin Paetz, The Fishes of Alberta, first published in 1970 and then revised in 1992, and especially the aforementioned four editions of Fishes of the World. His distinguished record of publications throughout the years resulted in Joe being selected, at the 2002 ASIH meetings in Kansas City, Missouri, as the recipient of the prestigious Robert H. Gibbs Award, presented by the society “for an outstanding body of published work in systematic ichthyology.”

Although the introduction of computers had by now greatly facilitated exchanges of information, the volume and complexity of new information made it necessary for the committee to meet, either yearly or every other year, for four-day work sessions, at which time ongoing issues and new information could be discussed and resolved on a face-to-face basis. Such meetings were normally held during the winter or early spring, and this, plus the fact that two of the committee members happened to reside in Gainesville, Florida, were factors in that city usually being selected as the meeting site. Claudine often used that opportunity to accompany Joe and escape the wintry weather of northern Canada for a few days.

Breaking with the usual pattern, the bi-yearly meeting of the c & sn of fishes committee was held in Mexico City in March 2009. At the meeting, Joe made a comment about feeling more fatigued than usual, although this was not immediately apparent to us and did not seem especially alarming, considering the somewhat thinner air associated with the higher elevation of Mexico City. At the ASIH meetings in Portland, Oregon, the following July, his fatigue had not abated, although it was not enough to prevent Joe from enjoying the meetings, which were highlighted by his election as ASIH President for the coming year.

Tests done shortly thereafter, however, showed the ongoing fatigue to be symptomatic of a serious medical problem, in which his bone marrow was not producing sufficient numbers of normal red and white blood cells. The condition, termed “Myelodysplastic Syndrome,” could progress to acute leukemia. This led to a regimen of blood transfusions and several experimental drugs, of which one seemed to hold the most promise for a prolonged and undiminished quality of life.

At the 2010 ASIH meetings in Providence, Rhode Island, Joe received the Robert K. Johnson Award for long-term distinguished service to the society. Although he was unable to attend the meetings, an arrangement was made by which members could communicate visually with Joe via the Internet. This allowed a large number of us to express our best wishes and hopes for improvement to his health.

In the meantime, it became evident that Joe's health would not allow him to fulfill his obligations as ASIH President-Elect, forcing him to resign this position, as he did also chairmanship of the committee on common and scientific names of fishes.

Despite this, the positive effects of his treatment during the subsequent year were sufficiently encouraging that Joe and Claudine were making plans to attend the 2011 ASIH meetings in Minneapolis, Minnesota, in July. A few weeks before the meeting, however, came the devastating news that the experimental drug was no longer working, and blood transfusions were the only option. From this point on, Joe's health underwent a precipitous decline, ending in his passing on August 9, 2011. In 2013, Joe's importance to ichthyology was recognized when ASIH established the Joseph S. Nelson Lifetime Achievement Award for an outstanding body of work in ichthyology.

At the 2012 ASIH meetings in Vancouver, British Columbia, Claudine presented a touching tribute to Joe during the plenary session. Her presentation brought home to us once again the realization that a wonderful friend and colleague was no longer with us. We will always miss you, Joe!


Carter Gilbert

Curator Emeritus

Florida Museum of Natural History

University of Florida

Gainesville, Florida

November 2015





Foreword


In 1976, Joseph S. Nelson published a modest volume with a bold title: Fishes of the World. Nelson was not the first to publish a book so named, but he was to become—for the next four decades—the only modern author associated with that title. This book, and the three editions that followed, are known simply by his last name: Nelson became the standard secondary reference for fish systematics.

To write Fishes of the World was a complex task. The first edition came 10 years after Greenwood et al.'s (1966) wholesale reorganization of fish classification, seven years after Gareth Nelson's (1969) classification of vertebrates based on the cladistic principles of Hennig (1966), and just three years after the audacious Interrelationships of Fishes of Greenwood, Miles, and Patterson (1973). For many, Nelson (1976) supplanted all of these because of its accessibility as a textbook and its philosophical aim: an eclectic mix of traditional and cladistic taxa diagnosed by both primitive and derived characters. A typical family account included the taxon, its common name, an outline sketch, a brief distribution, some maps (in the first two editions), a synopsis of characters (both primitive and derived), an estimate of the number of species, and some other comments that Joe considered to be of interest. Treatments could be uneven, yet this formula was repeated in all following editions. Nelson was cautious, especially about new taxa. His compromise was to discuss what he considered to be a radical proposal, such as that of the Neoteleostei by Rosen (1973), but not incorporate it into his classification.

After critical reviews of the first edition (e.g., Winterbottom, 1977, Journal of the Fisheries Research Board of Canada, 34:2431-2432), Joe published an expanded second edition in 1984. At 523 pages, the second Nelson (1984) was 107 pages longer than the first. Popularity of the first edition transferred to the second, and by the mid-1980s, before the age of the Internet, Joe Nelson and his book became a central clearing house for fish classification. Fish systematists enjoyed debating the latest classifications, and the largely morphological characters that supported them, with Joe at the annual meetings of the American Society of Ichthyologists and Herpetologists. Joe was kind and dignified as he engaged junior and senior ichthyologists with equal interest and respect. The newly published, monographic dissertations of an emerging generation—for example, Vari (1978) on terapontid perches, Johnson (1980) on lutjanids, Parenti (1981) on cyprinodontiforms, Stiassny (1981) on cichlids, and L. Grande (1982) on clupeomorphs—were cited and their classifications incorporated, in part, into Nelson (1984). This was validation of our place in the field of systematic ichthyology.

A decade passed between the second and third editions. Cladistic analysis and classification, once considered rogue, were now mainstream in systematic ichthyology. The pace of publication of revisionary studies of fishes increased, as did the third edition: Nelson (1994), at 600 pages, was the largest of the volumes to date. The distribution maps of the first two editions were eliminated due to space considerations. The third edition exposed more of Joe's unique views of characters and classification (Parenti, 1995, Copeia, 1: 262-264). He alone combined the billfish families Xiphiidae and Istiophoridae into the Xiphiidae. Further, he admitted to omitting characters that he found difficult to describe briefly, even if diagnostic. Many of the characters he listed are “field” or “key characters,” useful to identify specimens, but not necessarily to infer phylogenetic relationships. Joe absorbed criticism readily and alone. Despite the benefits a collaborator may have brought to the project, he never took on a co-author until asking Terry Grande and Mark Wilson to join him in writing the current edition, a job they would complete without him.

The fourth edition was published a decade ago. Dominance of the Internet meant that Nelson (2006) was no longer the sole, ready source of information on fish classification. The Catalog of Fishes, the spectacular achievement of Bill Eschmeyer, was available online (www.calacademy.org) and included fishes in a classification. Nonetheless, Nelson (2006) had become firmly established as the leading source of information on taxon diagnoses, summaries of species composition, and classification of fish taxa. Countless numbers of publications on fish systematics begin with a summary of number of species and other relevant data, citing the source as “Nelson (2006).” It stands alone as “the single most important book for fish classification” (Britz, 2006, Journal of Fish Biology, 69:1901).

The last edition of Fishes of the World that Joe wrote alone just hinted at the molecular phylogenetic revolution that was soon to come. These molecular studies and their proposed classificatory changes are summarized ably in this fifth edition by Grande and Wilson. Like Joe, they too use restraint in revising classifications and incorporate a judicious mix of the old and the new. There is a tension between using molecular versus morphological variation as phylogenetic data, and in what proportion, analogous to the tension that once existed between adopting a traditional versus a cladistic classification. Resolution of this tension is likely not imminent, and we may not wish it so, as it enlivens discussion and debate and moves systematic ichthyology onwards.


Lynne R. Parenti

Division of Fishes

Department of Vertebrate Zoology

National Museum of Natural History

Smithsonian Institution, Washington, DC

October 2015





Preface


Many years ago, when preparing to enter the American Museum of Natural History/CUNY doctoral program, I (TCG) was handed the first edition of Nelson's Fishes of the World by my Masters advisor who said, “Memorize it.” I spent that summer memorizing the contents of the book. My first day in Donn Rosen's ichthyology class, with book in hand, I was prepared—or at least I thought I was—and thus began my love of Fishes of the World. Little did I know that years later, I would receive a call from my friend and colleague Joe Nelson, to help him finish the fifth edition of the book. Life had come full circle and I was honored and ecstatic to work with Joe on this project. Sadly, my friend died soon after I agreed to help with the book, and my enthusiasm quickly turned to panic.

Mark Wilson (Joe's long-time colleague at the University of Alberta), had also agreed to help. We began sifting through Joe's library, files, and correspondence, trying to assemble the information needed to revise the book. We quickly realized that Joe had been unable to make much progress apart from gathering some scientific papers and accumulating comments and corrections from colleagues about the previous edition. We enlisted two amazing colleagues, W. Cal Borden (who was a postdoc in the Grande lab, and is now an Assistant Professor at Saginaw Valley State University) and John C. Bruner, research assistant in the Wilson lab at University of Alberta, to help search through the enormous volume of systematic and taxonomic publications that had appeared since 2005. We also later solicited the help of colleagues with expertise in specific fish groups to read and critique sections of the book (see Acknowledgments). However, any errors or omissions remain the responsibility of the authors.

Since the last edition of “Fishes” was published, the field of ichthyology has seen an explosion of published research, molecular systematics has become mainstream, new species descriptions have flourished, and large collaborative projects such as the NSF-funded Best Fish Phylogeny Project, the All Catfish Species Inventory Project, and the Cypriniform, Euteleost and Chondrichthyan Tree of Life projects have expanded our knowledge, challenged traditional paradigms, and stimulated new efforts. Transformative technologies in both morphology (e.g., 3-D scanning and imaging) and molecular systematics (e.g., Next-Gen Sequencing) are yielding their first results and promise to advance the field even further. In the paleontological and morphological realm, there have been international collaborations, meetings, and publications generated by the Early/Lower Vertebrates and the Mesozoic Fishes communities, as well as advanced anatomical, biomechanical, physiological, and ecological studies of extant fishes. This edition of Fishes of the World is written to reflect the present state of knowledge in the field, which is dynamic and constantly changing. This book shows how far we have come, and reflects current thinking, but by no means should it be taken as the final answer. There is much work still to do and there are many exciting discoveries ahead.

In the fifth edition of Fishes of the World, we have tried to preserve key features of the previous editions such as the iconic little fishes and the pithy and sometimes eccentric style of presentation that have been a hallmark of this much loved book. We also retain the style of scientific presentation for continuity with previous editions.

The fifth edition, however, differs in many respects from previous editions. We have tried to eliminate redundancy in the descriptions. To update diversity data such as numbers of species and genera, including synonymies, we used a combination of original literature and web-based resources. The systematic relationships and classifications of all fish groups were evaluated and updated in light of current, strongly supported research findings. In cases where there was conflict among studies and no convincing evidence for change, we retained the traditional placement of the group and discussed the conflict and alternative hypotheses. Systematic relationships are now represented by a greater number of simple cladograms. We have also added a significant number of new references and brought back important references dropped from previous editions.

Numerous additional fossil taxa, many of them discovered in recent years, have now been included. However, as in previous editions, space does not permit a comprehensive treatment of the diverse and remarkable fossil record of fishes.

Working on this edition has given us a renewed appreciation for the enormity of the task that Joe Nelson completed for each of the previous editions and a deeper understanding of the innovative and important work of our many ichthyological colleagues. It is our sincere hope that you, the reader, will find this edition of Fishes of the World to be worthy of the memory of Joe Nelson.





Acknowledgments


We wish to first thank the many people who have helped Joe throughout the years in the preparation of the first four editions of Fishes of the World. This edition builds on their efforts, and their help is greatly appreciated. Please see the complete list of acknowledgements in edition four, including the following: James Albert, M. Eric Anderson, Maria Elisabeth de Araújo, Gloria Arratia, William E. Bemis, Tim M. Berra, Jack C. Briggs, John C. Bruner, George H. Burgess, Kent E. Carpenter, Jeff C. Carrier, Marcelo de Carvalho, François Chapleau, Bruce C. Collette, Sara Collette, Leonard J. V. Compagno, Wilson J.E.M. Costa, Ed J. Crossman, Mrinal K. Das, Mario C. C. de Pinna, Dominique A. Didier, Brian S. Dyer, William N. Eschmeyer, Rainer Froese, Carter R. Gilbert, Nancy Gilbert, Lance Grande, Eileen D. Grogan, William C. Hamlett, David W. Greenfield, Harry J. Grier, Gavin F. Hanke, Carl L. Hubbs, Sir Ronald A. Javitch, Zerina Johanson, G. David Johnson, Maurice Kottelat, Dick Lund, John G. Lundberg, John G. Maisey, Keiichi Matsuura, Richard L. Mayden, John E. McCosker, Bob M. McDowall, John D. McEachran, Michal Miksik, Michael M. Mincarone, Masaki Miya, John F. Morrissey, Jack A. Musick, Heok Hee Ng, J. Ralph Nursall, Larry M. Page, Lynne R. Parenti, Nick V. Parin, Colin Patterson, Daniel Pauly, John R. Paxton, Ted W. Pietsch, E. Phil Pister, Francisco J. Poyato-Ariza, Jack E. Randall, Claude B. Renaud, Tyson R. Roberts, Ierecê L. Rosa, Richard H. Rosenblatt, Hans-Peter Schultze, Kwang-Tsao Shao, Stephen H. Shih, Gerald R. Smith, William F. Smith-Vaniz, Kenneth L. Soehn, Victor G. Springer, Melanie L. J. Stiassny, Hsi-Jen Tao, Bruce A. Thompson, Andrea Tintori, James C. Tyler, Edward O. Wiley.

We are grateful to the many colleagues who helped Joe via email correspondence during the early stages of revising this edition. We are aware of the following: Gloria Arratia, José Luís Birindelli, Alex Bannikov, Ralf Britz, Paulo Buckup, Meemann Chang, Kassi Cole, Bruce Collette, David Elliot, William Eschmeyer, Brett Human, G. David Johnson, Anne Kemp, Maurice Kottelat, Guillaume Lecointre, John Lundberg, Richard Mayden, John McCosker, Masaki Miya, Randall Mooi, Tom Near, Larry Page, Lynne Parenti, John Paxton, Leo Smith, Wayne Starnes, Jim Tyler, Peter Unmack, Cory Wakefield, Ed Wiley. We regret if we missed recognizing colleagues who corresponded with Joe in this regard. We are extremely grateful to all.

We wish to extend our deepest thanks to colleagues who have taken the time to offer suggestions for improvement and/or to read and critique specific sections of the book manuscript. Their expertise and extraordinary knowledge of fishes has greatly improved the published version: Gloria Arratia (Actinopterygii), Cal Borden (Lampriformes, Paracanthopterygii), John Bruner (Perciformes), Kerin Cleason (Batomorphi), Bruce Collette (Beloniformes, Istiophoriformes, Scombriformes), Kevin Conway (Cypriniformes and Gobiesociformes), Todd Cook (Elasmobranchii), Matt Davis (primitive Neoteleostei: Ateleopodomorpha, Cyclosquamata, Scopelomorpha), Dominique Didier (Holocephali), Christopher Fielitz (Aulopiformes), Katia Gonzalez-Rodriguez (primitive Actinopterygii), Lance Grande (basal crown Actinopterygii), Zeehan Jaafar (Gobiiformes), G. David Johnson(Elopomorpha, Ovalentaria), John Lundberg (Siluriphysi), John Maisey (Chondrichthyes), Jon Moore (Berycida), Lynne Parenti (Ovalentaria), Ted Pietsch (Lophiiformes), Leo Smith (Scorpaeniformes), James Tyler (Acanthuriformes and Tetradontiformes), Mark Westneat (Trachinoidei, Labroidei). Your advice was very important to us.

Special thanks go to Cal Borden, John Bruner and Todd Cook for the many hours they spent researching the literature, and for all of the fish discussion and debate that ensued.

We sincerely thank Carter Gilbert for contributing his remembrances of Joe, and Lynne Parenti for contributing the Foreword of the book and for all of her support throughout this process, especially during the last months.

I (TCG) wish to thank my EToL collaborators for five wonderful years of intense fish discussions, debate and learning together. My special thanks go to: Rich Broughton, Andrés Lópes, Guillermo Ortí, Leo Smith and Ed Wiley. If ever there was an academic guardian angel, it was Gloria Arratia. I wish to thank her for the many years of fish conversations, collaborations and friendship. Many thanks go also to John Maisey and Maria da Gloria P. de Carvalho for sharing their scientific expertise mixed with friendship, laughter and encouragement.

To all of our Mesozoic Fishes friends and colleagues we warmly thank you for the wonderful years of camaraderie, great fish discussions and insightful discovery. It is an honor to know you all. Ping pong, hiking in the woods (thanks H.-P.), and Mexican hot chocolate will always have special memories for us. Similarly, the many colleagues and friends who shared their expertise and congenial company during the meetings and field trips of the Early/Lower Vertebrates symposia and the Circum-Arctic Paleozoic Vertebrates project deserve special mention. I (MVHW) learned so much from all of you and felt right at home in your countries. Special thanks go to Tiiu Märss for teaching me so much about thelodonts and other agnathans. I also warmly thank my many colleagues at the University of Alberta who have taught me so much about biological diversity and earth history, with special thanks to Alison Murray in recent years for shared research experiences.

We thank Loyola University Chicago, Biology Department and the University of Alberta, Department of Biological Sciences for facilities and technical support in the preparation of this manuscript. Most of the fish figures from the previous edition were prepared by Pauly Wong (University of Alberta). Twenty-five new and replacement figures were prepared by Kofi Garbrah (University of Alberta) and a similar number by TCG. All cladograms and other diagrams were prepared by MVHW, who also produced artwork that became the basis for the front cover.

Thanks to the Wiley Group: Christopher Biray, Margaret Cummins, Michael New and Amy Odum for their professional assistance, patience, and encouragement.

The authors thank the Natural Sciences and Engineering Research Council of Canada for Discovery Grants to JSN (A5457), and MVHW (A9180) and the National Science Foundation for an AToL grant to TCG (DEB-0732589). This funding was invaluable in allowing us to conduct systematic research that was incorporated into this book.

Finally and most importantly, we wish to thank Joe Nelson for entrusting his book to us, and Claudine Nelson for giving us access to Joe's files and correspondence and for all of her support throughout the years. We are deeply grateful.


-T.C.G. & M.V.H.W.





Introduction


Over one-half—32,000 species and counting—of the world's living vertebrates—more than 60,000 species—are fishes. They arose and began to radiate more than 500 million years ago and both now and in the past exhibit incomparable diversity in their morphology, in the habitats they occupy, in their physiology, and in their behavior. This diversity is, in part, what makes understanding their evolutionary history and establishing a classification so challenging and yet fascinating. From hagfishes and lampreys to sharks, lungfishes and flatfishes, fishes include a vast array of amazing adaptations to almost all aquatic environments on earth.

Since the last edition of Fishes of the World (2006), a great many (thousands) of important studies on fish diversity, biology, morphology, and phylogenetic relationships have been published. We now have a much better understanding of their evolutionary relationships than we had even a decade ago. Science is a continually forward-moving search for knowledge, and this book reflects scientific knowledge about fishes as it exists today. As is normal in science, future researchers will build upon and improve upon what we know now.

The body of information known about fishes is vast and includes all aspects of biology. Fishes are fascinating to researchers because of the wealth of information and diversity to be found, both in fossil and living (extant) taxa. Since the 2006 version of this book, exciting new discoveries about fish morphology and evolution have been published. These include studies about the evolution of jaws, teeth, paired fins, internal fertilization, mimicry, hearing, and the biomechanics of feeding and locomotion. There have also been revolutionary findings concerning phylogenetic relationships, such as the hypotheses that extinct placoderms may be paraphyletic, that the Holostei are again monophyletic, that elopomorphs rather than osteoglossomorphs are the basal lineage of crown teleosts, that Lepidogalaxias is the most basal living euteleost, and that paracanthopterygians are once again united, though with revised membership.

The toolbox of the fish systematist has expanded to include molecular sequence analysis, evolutionary (including molecular) developmental biology, and technological advances such as 3D imaging to visualize complex internal morphology of both fossil and extant species. The new tools are being applied to many of what were the most intractable problems in fish systematics, including the relationships of rays, catfishes and percomorphs. There has also been an explosion in the number and variety of web-based databases and interactive tools, including Encyclopedia of Life, Phenoscape, Fishbase, Catalog of Fishes, Index to Organism Names, World Registry of Marine Species (WORMS), online academic libraries, journal publishers' web sites, the Paleobiology Database, and Fossilworks, all of which are available to almost any researcher, and all of which have been consulted by us. We also compiled and consulted thousands of original journal articles to better understand the current state of ichthyological knowledge.



Groups of fishes in which at least 20 new species were described in the decade 2005–2014. The area of each circle is approximately proportional to the number of new species in the group. The total number of new species described was about 3900, raising the total of known valid species to more than 32,000 (Eschmeyer and Fong, 2015).





Taxonomic Diversity


Since the 2006 version, more than 3890 species have been named. The species numbers of fishes given in the text, as in previous editions, are intended to be conservative estimates of valid described species, not of all named species nor of what might be undescribed. They are based, as far as possible, on the latest taxonomic revisions of families and genera and the opinions of the specialists.

There are 85 orders and 536 families of fishes recognized in this edition. Interestingly, about two-thirds of all species in the largest families are freshwater fishes, whereas only about 43% of all fishes are predominantly freshwater species. Freshwater habitats comprise only a small proportion of the earth's surface water, but contain a disproportionately large number of the world's fish species. The known diversity of both freshwater and marine fishes continues to increase rapidly, and the regions of the world where the greatest number of new discoveries are being made are indicated in the following figure.



Geographic regions of the world in which discoveries of new fish species have been most numerous. Newly discovered marine fishes have been found most often in the SW Caribbean, NW Indian Ocean, southern China Sea, and off the corners of Australia. Newly discovered freshwater fishes have been found most often in the Amazon Basin and the Parana Basin of South America, western Equatorial Africa, and southeastern Europe, especially near Turkey, as well as in Southeast Asia.





Importance to People


Fishes, like many other forms of life, are of immense value to humans. They have long been a staple item in the diet of many peoples, unfortunately leading to the downfall of many species (e.g., Atlantic Cod, Gadus morhua, fished to the brink of commercial extinction in the Western North Atlantic). Today fishes form an important element in the economy of many nations while giving incalculable recreational and psychological value to the naturalist, sports enthusiast, and home aquarist. They are also the subject of international and domestic agreements (Great Lakes Fishery Commission, a cross-border cooperation between the United States and Canada in the control of lamprey) and disagreements (suspension of albacore tuna fishing by Canadian fisherman in US waters in 2012). Many government institutions are devoted to the study of fish biology and propagation (e.g., propagation of Esox lucius and E. masquinongy for stocking by the Jake Wolfe Fish Hatchery, Illinois). Particular aspects of various species lend themselves to studies in behavior, ecology, evolution, genetics, and physiology. They are used as general indicators of pollution, partly to the direct benefit of humans and partly to protect what people consider a valuable and necessary part of their heritage and life.





Systematics and Classification


Systematics is the study of biological diversity, including reconstructing the phylogenetic (genealogical) relationships of organisms. Taxonomy is that part of systematics dealing with the theory and practice of describing diversity and erecting classifications. Classification is the practice of arranging items into groups or categories, and the resulting arrangement is called a classification. Taxa (singular taxon) are groups of organisms recognized in a classification and given biological names (e.g., Salmoniformes, Salmonidae, Oncorhynchus, Oncorhynchus nerka). A category is the level or rank at which the taxon is placed (e.g., order, family, genus, species). Generally, the objective in constructing a classification of a group of organisms is to reflect what are thought to be the evolutionary relationships of the various taxa in a hierarchical system of named groups.

We give examples of recognized generic names for each family; if the number is relatively small, we usually list them all. In choosing listed examples of generic names for large families, we have tried to choose: (1) genera with many species; (2) the type genus of the family, a subfamily, or a nominal family no longer recognized; (3) genera whose species exhibit some extreme biological diversity or unusual features, especially if mentioned in text; and (4) genera whose species are commonly encountered or are important in commercial fishery, sports fishery, or aquarium use. Generic synonyms are usually given only for genera recognized as valid in earlier editions of this book but that are now considered junior synonyms, or for cases when a family-group taxon is made synonymous because its type genus has been synonymized.

We consider fossils to be critical in understanding evolutionary relationships. Unfortunately, the fossil record in fishes is incomplete (more so in some than in other groups), and many decisions must be made with little or no evidence from fossils. However, we can answer many critical questions of interrelationships of higher taxa only with the assistance of the fossil record and not, conclusively at least, from extant material only. Many important fossil taxa are ranked along with extant taxa in the classification of this book, and many others are mentioned where appropriate.

A framework of formally named and ranked taxa is an important aid to understanding and communicating the implied relationships among groups of organisms. Completely unranked classifications are popular in some circles but are not used here because they communicate little information to those without specialized knowledge.

As in previous editions, we recognize a large number of named taxa in a formal hierarchy of taxonomic categories. The taxa are always intended to be monophyletic (i.e., clades), but in many cases future research may show that they are not, and the classification must be modified accordingly.

The categories used, and their endings in parentheses when consistent, are as follows: phylum, subphylum, superclass, grade, class, subclass, infraclass, division, subdivision, superorder (these 10 categories are centered in the text; the following categories are aligned left), series, subseries, infraseries, order (-iformes), suborder (-oidei), infraorder, superfamily (-oidea), family (-idae), subfamily (-inae), tribe (-ini), genus, subgenus, and species. Not all categories are employed within any one particular taxon. A dagger (†) denotes those taxa containing only fossil species.

Although there is a framework of named and ranked taxa, not all recognized (named) taxa are assigned a rank (i.e., placed in a named category). The following are examples of major taxa that are part of the classification but for which no formal rank is assigned: Vertebrata, Neoteleostei, and Acanthomorpha.

This edition, like earlier editions, adopts a simplified classification scheme, although even the simplified scheme can appear daunting. The number of categories and of named taxa are minimized by employing the “sequencing convention” for multiple named taxa at the same rank. For example, in a sequenced list of families within an order, the first family is the sister group of all others in the list, the second family is the sister of all except the first two, and so on. The last two families in the list are interchangeably each other's sisters. Consider this example classification of a hypothetical order with six families:

Order Numberiformes

Family Oneidae

Family Twoidae

Family Threeidae

Family Fouridae

Family Fiveidae

Family Sixidae



The sequence of the listed families tells us the phylogeny, using the sequencing convention. For example, Family Oneidae is the sister group of the clade of all five other families, and the Family Threeidae is the sister group of the clade of families Fouridae through Sixidae. The last two families could have been listed in either order, since they are each other's sisters. The tree that reflects these relationships is as follows:



In some larger taxa, we use the term “basal” or “most basal” when referring to the lineage, usually of low diversity, that is sister to all others in the group. Some call these more basal lineages “early-branching lineages.” Less often used but sometimes useful is the term “apical” or “more apical” to refer to a taxon that is high in the branching of the group's tree. A basal group is, of course, of equal age to its sister group and is not necessarily more primitive. In the tree shown above, the Family Oneidae would be considered basal while the families Fiveidae and Sixidae would be considered apical.

We report for many extant taxa the geologic age of the oldest reliably identified fossil members to give the reader an appreciation for the known fossil record. Such fossils give the minimum ages of lineages based on concrete data from the fossil record, even though in many cases the group in question must be considerably older than its first fossils because older fossils belonging to its sister-group are known. The difference between these two ages implies, for the lineage with the younger first fossil, a “ghost lineage”: a period of time during which it is inferred to have existed but is not recorded by fossils found to date.

It has become popular in phylogenetic literature to include estimates of past divergence times of lineages based on rates and amounts of molecular evolution. We do not list them or use them here.

In discussing fossil relationships, we make use of the concept of stem and crown groups. For any given taxon with extant members, the crown group is all those species descended from the last common ancestor (LCA) of all the extant members (see figure). Note that certain fossil taxa can be members of the crown group.



Tree diagram illustrating stem-group and crown-group concepts.

The stem group or stem-group taxa are all those extinct taxa known by fossils that are more closely related to this particular crown group than they are to any other extant clade. The Total Group is the sum of the stem group and the crown group. Using a real example, the Teleosteomorpha are the Total Group for teleostean fishes. The Teleocephala are the Crown Group teleosts.

A friendly word on the terms “fishes” and “fish” and on capitalizing their common names: The term “fishes” is properly used when referring to individuals of more than one species. However, when referring to one or more individuals of one species, the term “fish” is appropriate. Hence, it is correct to refer to 100 Rainbow Trout as fish, but to two different trouts, such as one Brook Trout and one Brown Trout, as fishes (the plural form Rainbow Trouts is discouraged). The common names of the three species given in this example (which happen to be in three different genera) were capitalized. The principles of common names in fishes established in 1960 by a joint committee of the American Fisheries Society and the American Society of Ichthyologists and Herpetologists, and explained in Nelson et al. (2004) are followed, except that the official common name of a species is treated as if it were a proper noun (see Nelson, Starnes, and Warren, 2002), as is common for some other groups of vertebrates (such as birds).





Anatomical Terminology


When given, the numbers of abdominal and caudal vertebrae are placed in parentheses after the total vertebral number—for example, 25 (10 + 15). When possible, the length is qualified by giving standard length (SL), fork length (FL), or total length (TL). Also included are estimated numbers of recognized (valid) genera and species (in some cases the number of species in each genus is also given). These figures are always for living forms, even if fossils are known for the taxon; selected fossil taxa are mentioned separately. The degree of agreement with these figures by specialists will vary from group to group (in part due to the subjective matter of lumping and splitting). For example, nearly everyone would agree that there are but two valid species of described percopsids, but one can easily find disagreement on the number of valid species of cichlids and gobiids that should be recognized.

Proposals to change the names of some bones from those used in previous editions to conform better with probable homologies have not been adopted unless otherwise indicated. For example, as noted in Janvier (1996) and Schultze (2008), what are commonly termed the frontals and parietals in actinopterygians, terms originally taken from human anatomy, are now known to be homologous with the parietals and postparietals, respectively, of early tetrapods.





Distribution and Biogeography


Fishes occur in lakes, streams, estuaries, and oceans throughout the world. In most species of fishes, all individuals live entirely either in fresh or in marine waters. Over 225 species are diadromous, regularly living part of their lives in lakes and rivers and part in the oceans. Among these, most are anadromous, spawning in fresh water but spending much of their time in the sea. A few are catadromous, spawning in the oceans but returning to fresh water. Classification of some species as marine, diadromous, estuarine, or freshwater is impossible, except as a generalization. Just as in an otherwise marine family there may be one species confined to fresh water, so in some species there are populations that occur in an environment opposite that of most others. Individuals of some otherwise marine species ascend rivers for short distances in part of their range, and those of some species that are usually freshwater are anadromous in some areas. Many freshwater and marine species are also common in brackish-water estuaries. About one-third of the 555 families have at least one species with individuals that spend at least part of their life in fresh water. Berra (2001) gives much information and distribution maps for the freshwater fish families.

Many environmental factors influence just where a certain species will predominate. Competition and other biological interactions may exert a strong influence along with physicochemical factors. In freshwater environments, species may show a preference for lakes or streams. Variations in preferences may exist over the range of a species. Among lakes they may show a preference for deep, cold, oligotrophic lakes or for shallower, warmer, and more productive mesotrophic and eutrophic lakes. In lake waters they may show a preference (horizontal and vertical) for the open-water limnetic zone, the benthic area, or shallow littoral areas. Fishes may even be restricted to certain types of bottom or do best under certain physicochemical conditions. Stream fishes may prefer riffle or quiet areas, and a zonation of species is usually found from the headwaters to the mouth. In the oceans, the vast majority of fishes are coastal or littoral. Most of those living beyond the 200-m-deep continental shelf (oceanic species) are deep-sea (mesopelagic, bathypelagic, abyssopelagic, or benthic at various depths); only a small minority regularly live close to the surface in the well-lighted upper 200-m zone (epipelagic), a region much larger in volume than the coastal waters. The epipelagic and mesopelagic fishes, which consist of both large predators and small plankton feeders, are varied, whereas most of the bathypelagic and abyssal fishes are relatively small.

Many species, both geologically young and old, have small ranges; the smallest is perhaps that of the Devils Hole Pupfish, Cyprinodon diabolis, found only in one spring in Ash Meadows, Nye County, Nevada. Many areas have a high degree of endemism. Marine fishes face the obvious land barriers (notably the New and Old World land masses) and mid-ocean barriers as well as many ecological and physiological barriers; freshwater species are limited by marine and land barriers. Some species have remarkably large ranges, and it would be interesting to know why some of their relatives have small ranges.

Over 130 marine species are known to extend around the world in tropical or subtropical waters. Many genera are represented in both the Pacific and Atlantic, but, almost always, different species are involved. Representatives of many marine genera and of some species occur in the temperate and polar faunas of both hemispheres. Individuals of some of these bipolar or antitropical taxa are surface-bound; others are deepwater. The vast majority of species, however, are tropical; most of the rest occur only in the Northern or only in the Southern Hemisphere. We know little of the abyssal depths and their species composition. Many abyssal species have been found at widely separated localities, which suggests that some may be virtually worldwide. No freshwater species is circumtropical, but two species, Esox lucius and Lota lota, are circumpolar and several others are almost so. No genus of freshwater fish has an antitropical distribution. Many freshwater fishes have shown a remarkable ability to disperse across newly exposed land areas following glaciation. In addition, they may occur in isolated waters in deserts as a result of a reduction of waters from times when drainage systems were connected.

In both fresh and marine waters, the largest number of species occurs in the tropics. There is a reduction toward the polar areas, although numbers of individuals in certain northern species are large. A great many species of freshwater fishes occur in tropical Africa, southeastern Asia, and the Amazon River—by far the world's largest river. For a tropical region, Central America has relatively few freshwater species because of the physiography and geological history of the area. Most oceanic islands lack indigenous fishes confined to fresh water, and continental areas recently exposed from the last ice age— for example, northern regions of North America, Europe (especially western Europe), and Asia—tend to have a relatively sparse fish fauna. In tropical areas, Africa exhibits the greatest diversity of nonostariophysan freshwater fishes; South America exhibits surprisingly little. In temperate areas, eastern North America shows the greatest diversity in nonostariophysan fishes. In marine waters, the Indo-West Pacific (Red Sea and Indian Ocean to northern Australia and Polynesia) is the richest, with the most species occurring in the New Guinea to Queensland area. In terms of diversity, southeastern Africa and Queensland appear to have the largest number of families of marine shorefishes. The West Indian or Caribbean fauna (southern Florida to northern Brazil) is also a rich one. The western African fauna, however, is relatively poor. Arctic and Antarctic faunas are depauperate. In all, the greatest number of fish species in the world inhabit the southeastern Asian region.



Broad surface thermal zones of the ocean, biogeographic regions of the continents, and native distribution of the family Cyprinidae, the most species-rich family of vertebrates. The biogeographical regions express degrees of endemism and are useful indicators of numbers and proportion of endemic organisms. We rarely use the continental regions in the text, and ichthyologists do not use them as much as in former times; the Nearctic and Palearctic are frequently combined into one region, the Holarctic. The thermal divisions of the sea denote tropical (or warm), subtropical, temperate, and cold (or polar) waters; warm temperate is sometimes used for all or part of the subtropical and warmer parts of the temperate (versus cool temperate) waters. Surface isotherms, used to define thermal regions, are subject to seasonal and annual changes. Major biogeographic regions recognized in the oceans include the Indo-West Pacific, tropical western Atlantic, tropical eastern Atlantic, North Pacific, North Atlantic, and Mediterranean-East Atlantic. Marine oceans share different similarities with one another; for example, for many families the tropical eastern Pacific shows a greater resemblance to the western Atlantic than to the Indo-West Pacific because of the mid-Pacific barrier and the relatively recent marine connection across the Isthmus of Panama. Information on the generalized thermal zones is based partly on Briggs (1974) and modified by numerous other sources. Distribution of the family Cyprinidae, shown by the shaded land area, is based on Berra (2001) and papers in Winfield and Nelson (1991).

Various methodological and philosophical approaches are used to explain the origin of distributional patterns of fishes, including areas of endemism. Both dispersal and vicariant events are important. Dispersal is regarded here as the movement, active or passive, of individuals to areas new to the existing population. Barriers of varying effectiveness may be involved as well as varying degrees of chance of reaching particular sites. It is of greatest biogeographic significance if the breeding range of the species is increased. Vicariance is the fragmentation of a former continuous distribution of the ancestral group into geographically separated units through the appearance of a barrier—for example, through plate tectonics. Both dispersal and vicariant approaches are used to explain disjunct distributions (the occurrence of a taxon in different areas with a marked geographical gap between them).

Examples of disjunct distributions include the following: occurrence of Prosopium coulteri in western North America and in Lake Superior; Geotria australis and Galaxias maculatus in Australia, New Zealand, and South America; cottids and agonids in cool temperate waters of the Northern and Southern Hemispheres; characiforms, aplocheiloids, and cichlids in Africa and South America. Plate tectonics had a profound effect on the distribution of many freshwater and marine fishes (e.g., it could well explain the occurrence of characiforms in South America and Africa), but not all disjunct distributions have a likely plate-tectonic explanation.





Human Impacts


We consider it desirable to maintain the fish diversity that systematists study, and systematists can play a leading role in protecting this diversity. We recognize the value of and our dependency upon fishes and other organisms, but our threats to the integrity of the environment also pose a serious threat to our fishes. As examples, according to the Great Barrier Reef Marine Park Authority, rising sea surface temperatures, ocean acidification, and rising sea level will negatively affect over 1600 species of fishes that live on the reef. Increased temperatures associated with lower dissolved oxygen levels in estuaries are affecting distribution patterns and abundance of Menhaden along the Atlantic and Gulf Coasts. The reduction of Menhaden, an important food source for larger fishes such as Bluefin (Pomatomus saltatix) and Bluefin Tuna (Thunnus thynnus), may likely have devastating effects on these fisheries.

Changing distribution patterns of many species and the extinction of native fishes has been directly linked to the human intervention. One recent and familiar example is the inadvertent introduction of the Round Goby (Neogobius melanostomus) native to central Eurasia, into the Great Lakes system via the ballast water of commercial tankers. In Lake Michigan, the Round Goby has outcompeted the native Mottled Sculpin (Cottus bairdi), causing its extirpation. The sheer mention of the famous Asian carp (actually the collective name for four species of carp that were imported in the 1970s to clean catfish ponds in the southern United States, but which then escaped and headed north) causes alarm. These carps are spreading widely in North American waterways, leading to habitat destruction and provoking sometimes unwise and expensive containment measures. Systematists have unique roles as experts on natural geographic distributions and as witnesses recording changes in biodiversity, whether natural or human-induced.





Phylum Chordata


Chordates are part of the superphylum Deuterostomia. The possible relationships of the chordates and deuterostomes to other metazoans were discussed by Halanych (2004). He restricted the deuterostomes to the chordates and their immediate sister group, comprising the hemichordates, echinoderms, and the wormlike Xenoturbella. Holland et al. (2015) reviewed the most popular theories about the origin of chordates.

The phylum Chordata has been used by most modern workers to encompass members of the subphyla Urochordata (tunicates or sea-squirts), Cephalochordata (lancelets), and Craniata (fishes, amphibians, reptiles, birds, and mammals). A fourth, fossil-only subphylum (†Conodontophorida, or conodonts) is also included here, although some authorities place it within the Craniata. Among extant taxa, the Cephalochordata and Craniata form a monophyletic group according to most authors (e.g., Cameron et al., 2000; Halanych, 2004) but some (e.g., Lowe et al., 2015) place Urochordata closest to Craniata.

Many exciting fossil finds in recent years reveal what the first fishes may have looked like, and these finds push the fossil record of fishes back into the early Cambrian, farther back than previously known. There is still much difference of opinion on the phylogenetic position of these new Cambrian species, and many new discoveries and changes in early fish systematics may be expected over the next decade. Some of the discoveries and controversies include the following.

As noted by Halanych (2004), Shu and collaborators have discovered fossil ascidian urochordates (e.g., †Cheungkongella), cephalochordate-like yunnanozoans (†Haikouella and †Yunnanozoon), and possible jawless craniates (†Myllokunmingia; its possible junior synonym is †Haikouichthys) over the last two decades, thus pushing the origins of these three major taxa at least back to the Early Cambrian (approximately 530–540 million years ago). These Early Cambrian fossils from Yunnan, China, lacked bone, but some of their soft anatomy was fossilized (Janvier, 1999; Shu et al., 1999).

Concerning †Myllokunmingia and †Haikouichthys, Shu et al. (1999) presented a phylogeny with †Myllokunmingia as sister to the remaining vertebrates and †Haikouichthys as sister to a clade with lampreys. Hou et al. (2002) described details of a new specimen co-occurring with the nominal †Myllokunmingia fengjiaoa and †Haikouichthys ercaicunensis and concluded that all are conspecific, favoring use of the oldest name †Myllokunmingia fengjiaoa. Characters include filamentous gills, V-shaped myomeres, and a distinct dorsal fin (the latter indicating a more derived condition than in the hagfish). Their phylogenetic analysis suggested that (the combined) †Myllokunmingia is either the sister group to the lampreys or the sister group to the vertebrates. However, not all authorities have agreed about the synonymy. Shu et al. (2003a) continued to argue that the taxa were distinct, placing †Haikouichthys either in a trichotomy with hagfishes and all vertebrates (i.e., including possibly as a stem craniate) or as the sister group to all vertebrates (i.e., in a position similar to that of †Myllokunmingia).

Concerning the yunnozoans, Shu et al. (2003b) and Shu and Morris (2003) proposed that the Early Cambrian †Haikouella and †Yunnanozoon are stem-group deuterostomes, and questionably placed them in the phylum †Vetulicolia, class †Yunnanozoa, family †Yunnanozoidae (= †Yunnanozoonidae). However, Mallatt et al. (2003) disagreed, interpreting †Haikouella not as a non-chordate stem-group deuterostome but rather as a chordate that is the immediate sister group of all craniates. The details of the relationships of these highly interesting fossils remain to be settled.

Other debates concern extant taxa, including a disagreement about the correct classification of the hagfishes (Myxiniformes) and the lampreys (Petromyzontiformes). This debate is discussed below under subphylum Craniata.

The following tree diagram illustrates the arrangement of the main chordate taxa adopted here and treated in the pages that follow. The diagram and the following pages include a selection of key fossil groups, each indicated by a shorter branch and by a dagger preceding its name.



Phylogeny of the main groups of Chordata. Explanation of symbols: †: taxon is extinct; *: taxon is paraphyletic according to some authorities.





SUBPHYLUM UROCHORDATA (Tunicata: the tunicates)


The tadpole larvae of tunicates possess gill slits, dorsal hollow nerve cord, notochord, and a muscular, unsegmented tail; the adults are usually sessile filter feeders and usually lack the preceding features. Feeding is by means of a mucous trap (i.e., endostyle) inside the pharynx as in cephalochordates and ammocoete larvae.

About 1,600 extant species are known, in three classes. Fossils include †Shankouclavis and possibly †Cheungkongella from the Early Cambrian Chengjiang biota of China (Shu et al., 2001; Chen et al., 2003). Although ascidian-like, their relationships to a particular class of Urochordata are unclear (Chen et al., 2003).





Class ASCIDIACEA (ascidians)


Larvae of ascidians free-swimming, tadpole-like (short-lived and nonfeeding); adults sessile benthic, solitary or colonial, and without a tail.

Ascidians are marine and distributed worldwide, extending from the intertidal well into the abyssal-benthic regions of the oceans.





Class THALIACEA (salps)


Larvae and adults of salps transparent, pelagic; adults may be solitary or colonial. They tend to be planktonic but are generally capable of weak movements. Remarkable life cycles are characteristic of this group, with sexual and asexual reproductive stages occurring.





Order PYROSOMIDA


Marine, worldwide except the Arctic Ocean. They can emit a strong phosphorescent light. Colonies tubular with a common atrial chamber and varying in length from about 3 cm to 1 m.





Order DOLIOLIDA (Cyclomyaria)


Marine; primarily tropical to temperate. Generally barrel-shaped with eight or nine muscle bands around the body.





Order SALPIDA (Hemimyaria)


Marine, all oceans. Cylindrical or prism-shaped.





Class APPENDICULARIA (Larvacea)


Pelagic, marine, all oceans from Arctic to Antarctic. Larval characteristics (such as tail) retained in adult.





SUBPHYLUM CEPHALOCHORDATA (Acrania, in part)


The notochord of cephalochordates extends to the anterior end of the body, anterior to the brain. Cranium absent; vertebrae absent; no cartilage or bone; heart consisting of a contractile vessel; red blood corpuscles absent; liver as a gut diverticulum; musculature segmented; epidermis as a single layer of cells; excretion by protonephridia with solenocytes; endostyle specialized (with iodine-fixing cells); true brain absent, but two pairs of cerebral lobes and nerves present; sexes separate.

About 30 species. Fossils likely include †Pikaia from the Middle Cambrian Burgess Shale of Canada, and †Cathaymyrus from the Early Cambrian Chengjiang fossil site in China (Shu et al., 1996; Conway Morris and Caron, 2012; Donoghue and Keating, 2014).

Cephalochordates and craniates share the following attributes (some also are present in the urocho