“…With the variety of both primitive and advanced teleosts living today, we are most emphatically of the opinion that approaches other than morphological ones would be exceedingly fruitful in the investigation of teleostean interrelationships” Greenwood et al. [1] .

New and Revised Classification for Bony Fishes based on Molecular Data (EToL)

The nomenclatural structure is based on the recent classification by Wiley and Johnson [2] using available morphological evidence gleamed from many investigators. Family names for bony fishes are based on Eschmeyer and Fong [3] and van der Laan et al. [4] , with minor modifications. Consult van der Laan et al. [4] for authorship of family names and Wiley and Johnson [10] for authorship of ordinal and subordinal names. This list is not intended as a comprehensive revision of valid family names; instead, it is simply an adaptation of their list based on published studies that we know validate or synonymize family groups using explicit phylogenetic evidence. Unlike Eschmeyer and Fong [3] and van der Laan et al. [4] , we do not recognize the family status of Anotopteridae, Omosudidae (synonyms of Alepisauridae [5] ) or Latidae (synonym of Centropomidae [6,7] ). Also, we recognize the following families, listed in Eschmeyer and Fong [3] and van der Laan et al. [4] as synonyms or subfamilies of other families: Botiidae (following [8] ), Diplophidae (following [9] ; apparently omitted by [3] ), Horabagridae (following [10] ), Sinipercidae (following [11] ), Steindachneriidae (following [12] ), Sudidae (following [5] ), and the pleuronectifoem Paralichthodidae, Poecilopsettidae, and Rhombosoleidae (following [13-15] ). A total of 498 families are recognized here, of which 366 (73.5%) were examined. Of these, 143 families included only one representative (39.2%) and 41 (18.4%) of the remaining 223 were rendered non-monophyletic in our analysis (non-monophyletic families are indicated below). For each order/suborder we list all families examined as well as the unexamined families whose taxonomic affinity is expected on the basis of traditional taxonomy or phylogenetic evidence. The list of unexamined families is also intended as a resource that may help fish systematists to direct future sequencing efforts.

New order-level or supraordinal taxa are erected on the basis of well-supported clades only (>90% bootstrap values). A total of 65 orders are recognized here, three of which appear to be new (Holocentriformes, Istiophoriformes, and Pempheriformes) and 16 resurrected or validated under a new circumscription. Some ordinal or subordinal names may appear to be new, but most are found in the literature at various hierarchical levels. As examples, Spariformes is a Bleeker name and Centrarchiformes is a Webber and de Beaufort name, both used as subfamily names. As priority is not applied to names above the family level, we have not made a thorough attempt to establish first use. Only those three for which no reference could be found are listed as “new.” New infraorders are named in Suborder Cottioidei to circumscribe well-corroborated clades and to conserve the rank of superfamily for subsequent revisions. The ordinal status of 55 percomorph families examined (as well as many others unexamined) belonging to Carangimorpharariae, Ovalentarariae, and Percomorpharia remains uncertain (i.e., incertae sedis) due to poor phylogenetic resolution. Percentages in parentheses following names indicate bootstrap support (no bootstrap values shown for redundant groups or monotypic taxa). The complete phylogenetic tree with annotated classification is shown here (OneZoom) or here (pdf).

Megaclass Osteichthyes (=Euteleostomi, =Euosteichthys)

Superclass Actinopterygii (100%)

Class Cladistia (100%)

Order Polypteriformes

Polypteridae

Class Actinopteri (100%)

Subclass Chondrostei (100%)

Order Acipenseriformes

Acipenseridae

Polyodontidae

Subclass Neopterygii (100%)

Infraclass Holostei (100%)

Order Amiiformes

Amiidae

Order Lepisosteiformes (100%)

Lepisosteidae

Infraclass Teleostei(100%)

Megacohort Elopocephalai sensu Arratia

[16] (100%)

Supercohort Elopocephala (100%)

Cohort Elopomorpha (100%)

Order Elopiformes (100%)

Megalopidae

Elopidae

Order Albuliformes (100%)

Albulidae

Order Notacanthiformes (100%)

Notacanthidae

Halosauridae

Order Anguilliformes (100%)

Eurypharyngidae

Saccopharyngidae

Anguillidae

Nemichthyidae

Muraenidae

Serrivomeridae

Congridae

Muraenesocidae

Ophichthidae

Not examined: Heterenchelyidae, Moringuidae, Chlopsidae, Myrocongridae, Synaphobranchidae, Colocongridae, Derichthyidae, Nettastomatidae, Cyematidae, Monognathidae, Protanguillidae.

Comment: Suborders recognized in Wiley and Johnson [2] based on previous work cited therein are significantly incongruent with the clades obtained in this analysis; thus, no subordinal classification is proposed.

Megacohort Osteoglossocephalai sensu Arratia [16] (38%)

Supercohort Osteoglossocephala (99%)

Cohort Osteoglossomorpha

Order Hiodontiformes (100%)

Hiodontidae

Order Osteoglossiformes (100%)

Pantodontidae

Osteoglossidae

Arapaimidae

Gymnarchidae

Mormyridae

Notopteridae

Supercohort Clupeocephala (100%)

Cohort Otomorpha, new circumscription (= Otocephala, Ostarioclupeomorpha) (100%)

Subcohort Clupei (87%)

Order Clupeiformes

Suborder Denticipitoidei

Denticipitidae

Suborder Clupeoidei (100%)

Clupeidae (not monophyletic)

Engraulidae

Chirocentridae

Pristigasteridae

Not examined: Dussumieriidae, Sundasalangidae

Subcohort Alepocephali (100%)

Order Alepocephaliformes

Alepocephalidae (not monophyletic)

Platytroctidae

Not examined: Leptochilichthyidae

Subcohort Ostariophysi (100%)

Section Anotophysa (97%)

Order Gonorynchiformes

Suborder Gonorynchoidei

Gonorynchidae

Suborder Chanoidei

Chanidae

Suborder Knerioidei (100%)

Kneriidae

Not examined: Phractolaemidae

Section Otophysa (100%)

Superorder Cyprinae (100%)

Order Cypriniformes

Gyrinocheilidae

Catostomidae

Botiidae

Nemacheilidae

Cobitidae

Cyprinidae

Not examined: Psilorhynchidae, Balitoridae, Vaillantellidae

Superorder Characiphysae (100%)

Order Gymnotiformes (100%)

Gymnotoidei (not monophyletic)

Gymnotidae (not monophyletic)

Apteronotidae

Sternopygidae (not monophyletic)

Rhamphichthyidae

Not examined: Hypopomidae

Comment: Relationships recovered here are incongruent with the subordinal classification in Wiley and Johnson [2] based on Albert [17] and thus no subordinal classification is proposed.

Order Characiformes (100%)

Suborder Citharinoidei (not monophyletic)

Distichodontidae

Citharinidae

Suborder Characoidei (not monophyletic)

Chilodontidae

Ctenoluciidae

Crenuchidae

Alestidae

Hepsetidae

Erythrinidae

Cynodontidae

Serrasalmidae

Parodontidae

Hemiodontidae

Curimatidae

Prochilodontidae

Lebiasinidae

Chalceidae

Bryconidae

Triportheidae

Characidae

Gasteropelecidae

Acestrorhynchidae

Not examined: Anostomidae, Iguanodectidae

Comment: although not monophyletic in this analysis, the monophyly of characiform suborders has been corroborated by other molecular studies (e.g., [18]).

Order Siluriformes (100%)

Suborder Loricaroidei (93%)

Nematogenyidae

Trichomycteridae

Loricariidae

Astroblepidae

Callichthyidae

Suborder Diplomystoidei

Diplomystidae

Suborder Siluroidei (100%)

Sisoridae

Akysidae

Amblycipitidae

Bagridae

Schilbeidae

Horabagridae

Heteropneustidae

Clariidae

Chacidae

Cetopsidae

Cranoglanididae

Ictaluridae

Pangasiidae

Anchariidae

Ariidae

Claroteidae

Amphiliidae

Malapteruridae

Mochokidae

Siluridae

Plotosidae

Doradidae

Auchenipteridae

Heptapteridae

Pseudopimelodidae

Pimelodidae

Not examined: Austroglanididae, Erethistidae, Aspredinidae. Lacantuniidae, Olyridae, Scoloplacidae.

Supercohort Clupeocephala (cont.)

Cohort Euteleosteomorpha (100%)

Subcohort Lepidogalaxii

Order Lepidogalaxiiformes

Lepidogalaxiidae

Subcohort Protacanthopterygii sedis mutabilis (37%)

Order Galaxiiformes, new circumscription (100%)

Galaxiidae

Order Argentiniformes, new circumscription (100%)

Bathylagidae

Microstomatidae

Opisthoproctidae

Argentinidae

Order Salmoniformes (100%)

Salmonidae

Order Esociformes (100%)

Umbridae

Esocidae

Subcohort Stomiatii, new circumscription (73%)

Order Stomiatiformes (100%)

Diplophidae

Gonostomatidae

Phosichthyidae (not monophyletic)

Stomiidae (not monophyletic)

Sternoptychidae

Order Osmeriformes, new circumscription (100%)

Retropinnidae

Osmeridae

Plecoglossidae

Salangidae

Subcohort Neoteleostei, new circumscription (100%)

Infracohort Ateleopodia (100%)

Order Ateleopodiformes

Ateleopodidae

Infracohort Eurypterygia (96%)

Section Aulopa (100%)

Order Aulopiformes

Suborder Aulopoidei (not monophyletic)

Synodontidae (not monophyletic)

Aulopidae

Pseudotrichonotidae

Suborder Paraulopoidei

Paraulopidae

Suborder Alepisauroidei (not monophyletic)

Giganturidae

Bathysauridae

Ipnopidae

Scopelarchidae (not monophyletic)

Notosudidae

Chlorophthalmidae (not monophyletic)

Paralepididae (not monophyletic)

Evermannellidae

Sudidae

Alepisauridae

Not examined: Bathysauroididae, Bathysauropsidae

Comment: Suborders and families listed are as in Davis [5]. Although not monophyletic herein, the monophyly aulopiform suborders was supported by Davis [5].

Section Ctenosquamata (97%)

Subsection Myctophata (100%)

Order Myctophiformes

Neoscopelidae

Myctophidae

Subsection Acanthomorphata (97%)

Division Lampridacea (100%)

Order Lampridiformes, new circumscription

Lampridae

Lophotidae

Regalecidae

Trachipteridae

Not examined: Veliferidae, Radiicephalidae.

Division Paracanthomorphacea sensu Grande et al. (in press) (93%)

Series Percopsaria (100%)

Order Percopsiformes

Percopsidae

Aphredoderidae

Amblyopsidae

Series Zeiogadaria(=Zeioigadiformes sensu Li et al. [19]) (98%)

Subseries Zeariae (100%)

Order Zeiformes

Zeidae

Parazenidae

Not examined: Cyttidae, Oreosomatidae, Zenionidae, Grammicolepididae.

Subseries Gadariae (100%)

Order Stylephoriformes (sensu Miya et al. [20])

Stylephoridae

Order Gadiformes (100%)

Macrouridae (not monophyletic)

Steindachneriidae

Merlucciidae

Moridae

Phycidae

Lotidae

Gadidae

Not examined: Muraenolepididae, Euclichthyidae, Bregmacerotidae, Melanonidae,

Comments: The monophyly of several gadiform suborders sensu Roa-Varon and Orti [12] was not recovered, therefore we do not provide a subordinal classification.

Division Polymixiacea (100%)

Order Polymixiiformes

Polymixiidae

Division Euacanthomorphacea sensu [21] (99%)

Subdivision Berycimorphaceae (87%)

Order Beryciformes, new circumscription (circumscription similar to Trachichthyiformes sensu Moore [22])

Diretmidae

Monocentridae

Trachichthyidae

Anoplogastridae

Berycidae

Melamphaidae

Barbourisiidae

Rondeletiidae

Cetomimidae

Not Examined: Anomalopidae, Gibberichthyidae, Hispidoberycidae, Stephanoberycidae (placement expected following Moore [22]).

Subdivision Holocentrimorphaceae (100%)

Order Holocentriformes, new

Holocentridae

Comments: and Moore [22],Stiassny and Moore [23] provide morphological evidence supporting a sister-group relationship between holocentrids and percomorphs, which further supports placement of this family in its own order.

Subdivision Percomorphaceae

Subdivision Percomorphaceae (cont.)

Not Examined (traditionally placed in “Perciformes”): Aplodactylidae, Banjosidae, Bathyclupeidae, Callanthiidae, Centracanthidae, Cepolidae, Champsodontidae, Chironemidae, Datnioididae, Dichistiidae, Dinolestidae, Dinopercidae, Hapalogenyidae, Lactariidae, Latridae, Ostracoberycidae, Parascorpididae, Pentacerotidae, Perciliidae, Scombropidae, Symphysanodontidae, Trichodontidae, Trichonotidae.

Series Ophidiimorpharia (100%)

Order Ophidiiformes

Suborder Ophidioidei

Ophidiidae

Not examined: Carapidae

Suborder Bythitoidei

Bythitidae

Not examined: Aphyonidae, Parabrotulidae

Series Batrachoidimorpharia (100%)

Order Batrachoidiformes

Batrachoididae

Series Gobiomorpharia (see also [24,25]) (100%)

Order Kurtiformes, new circumscription (98%)

Kurtidae

Apogonidae

Comments: Johnson [26] noted that the configuration of the dorsal gill-arch elements may be homologous in Kurtus and apogonids.

Order Gobiiformes (100%)

Suborder Odontobutoidei (100%)

Odontobutidae

Suborder Eleotroidei (97%)

Eleotridae

Suborder Gobioidei (100%)

Gobiidae (not monophyletic)

Microdesmidae

Not examined: Rhyacichthyidae, Kraemeriidae, Schindleriidae, Thalasseleotrididae, Xenisthmidae.

Series Scombrimorpharia (97%)

Order Syngnathiformes, new circumscription (97%)

Syngnathidae (not monophyletic)

Fistulariidae

Centriscidae

Aulostomidae

Dactylopteridae

Creediidae

Callionymidae

Mullidae

Not examined: Pegasidae, Solenostomidae, Draconettidae (assumed affinity with Callionymidae), Leptoscopidae (assumed affinity with Creediidae [27]).

Order Scombriformes, new circumscription (=Stromateoidei sensu Li et al. [19]) (100%)

Centrolophidae

Nomeidae

Ariommatidae

Chiasmodontidae

Gempylidae

Trichiuridae

Icosteidae

Stromateidae

Bramidae

Scombridae (not monophyletic)

Pomatomidae

Caristiidae

Scombrolabracidae

“Stromateoid” families not examined: Amarsipidae, Tetragonuridae; not examined but assumed affinity: Arripidae [28]

Series Carangimorpharia (94%)

Subseries Anabantomorpharariae (=Anabantiformes sensu Li et al. [19]) (99%)

Order Synbranchiformes, new circumscription (98%)

Suborder Indostomoidei

Indostomidae

Suborder Synbranchoidei

Synbranchidae

Suborder Mastacembeloidei

Mastacembelidae

Not examined: Chaudhuriidae

Order Anabantiformes (100%)

Suborder Anabantoidei (95%)

Osphronemidae

Helostomatidae

Anabantidae

Not examined: Badidae, Pristolepididae

Suborder Channoidei (85%)

Channidae

Nandidae

Subseries Carangimorpharariae (=Carangimorpha sensu Li et al. [19]) (100%)

Incertae sedis: Centropomidae (sensu Greenwood), Sphyraenidae, Toxotidae, Leptobramidae, Menidae, Polynemidae.

Order Istiophoriformes, new (100%)

Istiophoridae

Xiphiidae

Order Carangiformes sedis mutabilis (not monophyletic)

Carangidae

Echeneidae

Coryphaenidae

Rachycentridae

Nematistiidae

Comment: Although is rendered non-monophyletic, monophyly of the order is not significantly rejected (see discussion in [15]).

Order Pleuronectiformes sedis mutabilis (not monophyletic)

Suborder Psettodoidei (100%)

Psettodidae

Suborder Pleuronectoidei (100%)

Achiridae

Achiropsettidae

Bothidae

Citharidae

Cynoglossidae

Paralichthyidae (not monophyletic)

Pleuronectidae

Poecilopsettidae

Rhombosoleidae (not monophyletic)

Samaridae

Scophthalmidae

Soleidae

Not examined: Paralichthodidae

Comment: Although Psettodidae was not the sister group of pleuronectoids in the analysis, the order was resolved as monophyletic by a recent study, albeit with low support [15].

Subseries Ovalentarariae (sensu Smith & Near in Wainwright et al. [29]; = Stiassnyiformes sensu Li et al. [19]) (100%)

Incertae sedis: Embiotocidae, Grammatidae (in part; not monophyletic), Plesiopidae, Polycentridae, Pomacentridae, Pseudochromidae (not monophyletic), Ambassidae.

Superorder Cichlomorphae

Order Cichliformes, new circumscription (93%)

Cichlidae

Pholidichthyidae

Superoder Atherinomorphae (100%)

Order Atheriniformes (100%)

Atherinopsidae

Atherinidae

Melanotaeniidae

Bedotiidae

Pseudomugilidae

Telmatherinidae

Notocheiridae

Not examined: Phallostethidae, Isonidae, Dentatherinidae.

Order Beloniformes (32%)

Suborder Adrianichthyoidei

Adrianichthyidae

Suborder Exocoetoidei (100%)

Belonidae (not monophyletic)

Exocoetidae (not monophyletic)

Zenarchopteridae (not monophyletic)

Hemiramphidae (not monophyletic)

Scomberesocidae

Order Cyprinodontiformes (57%)

Suborder Aplocheiloidei

Aplocheilidae

Suborder Cyprinodontoidei (100%)

Cyprinodontidae

Poeciliidae

Fundulidae

Not examined: Rivulidae, Profundulidae, Goodeidae, Valenciidae, Anablepidae, Nothobranchiidae.

Superorder Mugilomorphae (100%)

Order Mugiliformes

Mugilidae

Superorder Blenniimorphae (100%)

Incertae sedis: Grammatidae (in part; not monophyletic) and Opistognathidae.

Order Blenniiformes, new circumscription (Li et al. [19] plus Gobiesocidae) (100%)

Gobiesocidae

Blenniidae

Dactyloscopidae

Labrisomidae (not monophyletic)

Clinidae

Chaenopsidae

Tripterygiidae

Series Percomorpharia (99%)

Incertae sedis: Acropomatidae (not monophyletic), Caesionidae, Caproidae, Centrogenyidae, Chaetodontidae, Emmelichthyidae, Enoplosidae, Epigonidae, Gerreidae, Haemulidae, Howellidae, Kuhliidae, Kyphosidae (not monophyletic), Lateolabracidae, Leiognathidae, Lobotidae, Lutjanidae (not monophyletic), Malacanthidae, Monodactylidae, Moronidae, Oplegnathidae, Percichthyidae (not monophyletic), Polyprionidae (not monophyletic), Pomacanthidae, Priacanthidae, Scatophagidae, Sciaenidae, Siganidae, Sillaginidae, Sinipercidae, Terapontidae.

Order Uranoscopiformes, new circumscription (= Paratrachiniformes sensu Li et al. [19]) (95%)

Uranoscopidae

Ammodytidae

Pinguipedidae

Not examined: Cheimarrichthyidae

Order Labriformes sensu stricto (100%)

Odacidae

Labridae (not monophyletic)

Scaridae

Order Ephippiformes, new circumscription (100%)

Drepaneidae

Ephippidae

Comments: Greenwood et al. [1] hypothesized a close affinity between Drepane and ephippids.

Order Spariformes sensu Akazaki [30] (94%)

Lethrinidae

Sparidae

Possibly included (examined): Nemipteridae

Comments: Akazaki [30] proposed these families to be closely related based on specializations of the suspensorium and other features [26].

Order Lophiiformes (100%)

Suborder Lophioidei (100%)

Lophiidae

Suborder Antennarioidei (100%)

Antennariidae

Not examined: Tetrabrachiidae, Lophichthyidae, Brachionichthyidae

Suborder Chaunacoidei (100%)

Chaunacidae

Suborder Ogcocephaloidei (100%)

Ogcocephalidae

Suborder Ceratioidei (not monophyletic) (87%)

Ceratiidae (not monophyletic)

Gigantactinidae

Himantolophidae

Melanocetidae

Oneirodidae

Not examined: Caulophrynidae, Neoceratiidae, Diceratiidae, Thaumatichthyidae, Centrophrynidae, Linophrynidae.

Order Tetraodontiformes (100%)

Suborder Triacanthodoidei (100%)

Triacanthodidae

Suborder Tetraodontoidei, new circumscription (100%)

Diodontidae

Tetraodontidae

Suborder Moloidei, new

Molidae

Suborder Balistoidei, new circumscription (100%)

Monacanthidae

Balistidae

Suborder Ostracioidei, new (100%)

Ostraciidae

Aracanidae

Suborder Triacanthoidei, new

Triacanthidae

Not examined or classified: Triodontidae

Comment: suborders recognized by Santini and Tyler [31] are not recovered here or in previous molecular studies and thus a new subordinal classification is proposed.

Order Acanthuriformes, restricted circumscription (see also [32]) (100%)

Luvaridae

Zanclidae

Acanthuridae

Order Pempheriformes, new (100%)

Glaucosomatidae

Pempheridae

Comments: Tominaga [33] suggested that features of the cranium and swimbladder may be homologous in Pempheris and Glaucosoma.

Order Cirrhitiformes, new circumscription (91%)

Cheilodactylidae

Cirrhitidae

Order Centrarchiformes, new circumscription (100%)

Centrarchidae

Elassomatidae

Order Perciformes, new circumscription (89%)

Incertae sedis: Serranidae, Platycephalidae

Not examined (traditionally placed in Scorpaeniformes): Apistidae, Aploactinidae, Bembridae, Congiopodidae, Cottocomephoridae, Eschmeyeridae, Gnathanacanthidae, Hoplichthyidae, Neosebastidae, Pataecidae, Plectrogeniidae, Setarchidae, Synanceiidae, Tetrarogidae, Zanclorhynchidae.

Suborder Percoidei, restricted circumscription (100%)

Percidae

Not examined: Trachinidae.

Comment: Lautredou et al. [34] using seven nuclear markers obtained a clade uniting Percidae and Trachinidae with full support.

Suborder Notothenioidei (100%)

Pseudaphritidae

Eleginopsidae

Nototheniidae (not monophyletic)

Artedidraconidae (not monophyletic)

Harpagiferidae

Bathydraconidae (not monophyletic)

Channichthyidae

Bovichtidae

Suborder Scorpaenoidei, new circumscription (83%)

Scorpaenidae (not monophyletic)

Sebastidae (not monophyletic)

Suborder Triglioidei sensu Jordan [35] (100%)

Triglidae

Peristediidae

Suborder Cottioidei, new circumscription (= Cottimorpha sensu sensu Li et al. [19]) (100%)

Comment: We have chosen to recognize clades within this suborder as infraorders, adopting the ending –ales for this rank.

Infraorder Anoplopomatales, new

Anoplopomatidae

Infraorder Gasterosteales (100%)

Gasterosteidae

Hypoptychidae

Aulorhynchidae (not monophyletic)

Infraorder Cottales (96%)

Hexagrammidae (not monophyletic)

Cyclopteridae

Liparidae

Cottidae (not monophyletic)

Agonidae (not monophyletic)

Psychrolutidae

Not examined, traditionally placed in Cottoidei: Rhamphocottidae, Ereuniidae, Hemitripteridae, Abyssocottidae, Comephoridae, Bathylutichthyidae, Normanichthyidae, Parabembridae.

Infraorder Zoarcales (100%)

Bathymasteridae (not monophyletic)

Stichaeidae (not monophyletic)

Zaproridae

Cryptacanthodidae

Pholidae

Anarhichadidae

Zoarcidae

Not examined: Ptilichthyidae, Scytalinidae.

Superclass Sarcopterygii (96%)

Class Coelacanthimorpha (= Actinistia)

Order Coelacanthiformes