Appendix 2 from Betancur et al 2013
Revised Classification for Bony Fishes (v.1)
Version Date: 18 April 2013 -- this version has been superseded by version 2 and then by version 3 on 31 July 2014
The nomenclatural arrangement presented here builds on the existing classification by Wiley and Johnson [1] and intends to preserve names and taxonomic composition of groups whenever possible. However, adjustments are made to recognize new well-supported molecular clades, many of which also have been obtained by previous molecular studies (several examples discussed below). Order-level or supraordinal taxa are erected (new) or resurrected on the basis of well-supported clades only (>90% bootstrap values). Current taxon names supported by previous molecular or morphological studies are retained if congruent with our results, even if bootstrap support is low (e.g., Osteoglossocephalai sensu Arratia [2] with only 38% bootstrap). In some cases, ordinal or subordinal taxa that were not monophyletic in our analysis are also validated, as long as the incongruence is not supported by strong bootstrap values. Examples include the suborder Blennioidei (not monophyletic here but monophyletic in Wainwright et al. [3]) and the order Pleuronectiformes (not monophyletic here but monophyletic in Betancur-R. et al. [4]).
Family names for bony fishes are based on Eschmeyer and Fong [5] and van der Laan et al. [6], with minor modifications. Consult van der Laan et al. [6] for authorship of family names and Wiley and Johnson [10] for authorship of ordinal and subordinal names. Our 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 [5] and van der Laan et al. [6], we do not recognize the family status of Anotopteridae, Omosudidae (synonyms of Alepisauridae [7]) or Latidae (synonym of Centropomidae [8,9]). Also, we recognize the following families, listed in Eschmeyer and Fong [5] and van der Laan et al. [6] as synonyms or subfamilies of other families: Botiidae (following [10]), Diplophidae (following Nelson [11]; apparently omitted by Eschmeyer and Fong [5]), Horabagridae (following Sullivan et al. [12]), Sinipercidae (following Li et al. [13]), Steindachneriidae (following Roa-Varon and Orti [14]), Zanclorhynchidae, the aulopiform Bathysauropsidae and Sudidae (following Davis [7]), and the pleuronectiform Paralichthodidae, Poecilopsettidae, and Rhombosoleidae (following Chapleau [15], Munroe [16]). A total of 502 families are recognized here, of which 369 (73.5%) were examined. Of these, 146 families included only one representative (39.6%) and 40 (17.9%) 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.
A total of 66 orders are classified, three of which are new (Holocentriformes, Istiophoriformes, and Pempheriformes), and 15 are resurrected or validated under a new circumscription. Some ordinal or subordinal names may appear to be new, but most can be found in the literature at various hierarchical levels. As examples, Spariformes is a Bleeker name and Centrarchiformes is a Webber and de Beaufort name. 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 may conserve the rank of superfamily in subsequent revisions. The ordinal status of 50 percomorph families examined (as well as many others unexamined) belonging to Carangimorphariae, Ovalentariae, 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 illustrated in Fig. S1. The new classification scheme presented here should be considered a work in progress (version 1), as any other hypothesis. It is likely to include involuntary errors and omissions in addition to the many unexamined, sedis mutabilis, and incertae sedis taxa. Updates should be forthcoming as new evidence become available and feedback from experts help refine it. For the most updated version visit DeepFin.
Megaclass Osteichthyes (=Euteleostomi, =Euosteichthyes)
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 [2] (100%)
Supercohort Elopocephala (100%)
Cohort Elopomorpha (100%)
Order Elopiformes (100%)
Elopidae
Megalopidae
Order Albuliformes (100%)
Albulidae
Order Notacanthiformes (100%)
Halosauridae
Notacanthidae
Order Anguilliformes (100%)
Anguillidae
Congridae
Eurypharyngidae
Muraenesocidae
Muraenidae
Nemichthyidae
Ophichthidae
Saccopharyngidae
Serrivomeridae
Not examined: Chlopsidae, Colocongridae, Cyematidae, Derichthyidae, Heterenchelyidae, Monognathidae, Moringuidae, Myrocongridae, Nettastomatidae, Protanguillidae, Synaphobranchidae.
Comment: Suborders recognized in Wiley and Johnson [1] 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 [2] (38%)
Supercohort Osteoglossocephala sensu Arratia [2] (99%)
Cohort Osteoglossomorpha
Order Hiodontiformes (100%)
Hiodontidae
Order Osteoglossiformes (100%)
Arapaimidae
Gymnarchidae
Mormyridae
Notopteridae
Osteoglossidae
Pantodontidae
Supercohort Clupeocephala sensu Arratia [17] (100%)
Cohort Otomorpha, new circumscription (=Otocephala, Ostarioclupeomorpha) (100%)
Subcohort Clupei (87%)
Order Clupeiformes
Suborder Denticipitoidei
Denticipitidae
Suborder Clupeoidei (100%)
Chirocentridae
Clupeidae (not monophyletic)
Engraulidae
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
Not examined: Phractolaemidae.
Suborder Chanoidei
Chanidae
Suborder Knerioidei (100%)
Kneriidae
Section Otophysa (100%)
Superorder Cyprinae (100%)
Order Cypriniformes
Botiidae
Catostomidae
Cobitidae
Cyprinidae
Gyrinocheilidae
Nemacheilidae
Not examined: Balitoridae, Barbuccidae, Ellopostomatidae, Psilorhynchidae, Serpenticobitidae, Vaillantellidae.
Superorder Characiphysae (100%)
Order Gymnotiformes (100%)
Suborder Gymnotoidei (not monophyletic)
Gymnotidae (not monophyletic)
Suborder Sternopygoidei (not monophyletic)
Apteronotidae
Rhamphichthyidae
Sternopygidae (not monophyletic)
Not examined: Hypopomidae.
Comment: Although not monophyletic here, the monophyly gymnotiform suborders was corroborated by Albert and Crampton [18].
Order Characiformes (100%)
Suborder Citharinoidei (not monophyletic)
Citharinidae
Distichodontidae
Suborder Characoidei (not monophyletic)
Acestrorhynchidae
Alestidae
Bryconidae
Chalceidae
Characidae
Chilodontidae
Crenuchidae
Ctenoluciidae
Cynodontidae
Erythrinidae
Gasteropelecidae
Hemiodontidae
Hepsetidae
Lebiasinidae
Parodontidae
Prochilodontidae
Serrasalmidae
Triportheidae
Not examined: Anostomidae, Curimatidae, Iguanodectidae.
Comment: Although not monophyletic in this analysis, the monophyly of characiform suborders has been corroborated by other molecular studies (e.g., [19]).
Order Siluriformes (100%)
Suborder Loricaroidei (93%)
Astroblepidae
Callichthyidae
Loricariidae
Nematogenyidae
Trichomycteridae
Suborder Diplomystoidei
Diplomystidae
Suborder Siluroidei (100%)
Akysidae
Amblycipitidae
Amphiliidae
Anchariidae
Ariidae
Auchenipteridae
Bagridae
Cetopsidae
Chacidae
Clariidae
Claroteidae
Cranoglanididae
Doradidae
Heptapteridae
Heteropneustidae
Horabagridae
Ictaluridae
Malapteruridae
Mochokidae
Pangasiidae
Pimelodidae
Plotosidae
Pseudopimelodidae
Schilbeidae
Siluridae
Sisoridae
Not examined: Aspredinidae, Austroglanididae, Erethistidae, 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%)
Argentinidae
Bathylagidae
Microstomatidae
Opisthoproctidae
Order Salmoniformes (100%)
Salmonidae
Order Esociformes (100%)
Esocidae
Umbridae
Subcohort Stomiatii, new circumscription (73%)
Order Stomiatiformes (=Stomiiformes) (100%)
Diplophidae
Gonostomatidae
Phosichthyidae (not monophyletic)
Sternoptychidae
Stomiidae (not monophyletic)
Order Osmeriformes, new circumscription (100%)
Osmeridae
Plecoglossidae
Retropinnidae
Salangidae
Subcohort Neoteleostei, new circumscription (100%)
Infracohort Ateleopodia (100%)
Order Ateleopodiformes
Ateleopodidae
Infracohort Eurypterygia (96%)
Section Aulopa (100%)
Order Aulopiformes
Suborder Aulopoidei (not monophyletic)
Aulopidae
Pseudotrichonotidae
Synodontidae (not monophyletic)
Suborder Paraulopoidei
Paraulopidae
Suborder Alepisauroidei (not monophyletic)
Alepisauridae
Bathysauridae
Chlorophthalmidae (not monophyletic)
Evermannellidae
Giganturidae
Ipnopidae
Notosudidae
Paralepididae (not monophyletic)
Scopelarchidae (not monophyletic)
Sudidae
Not examined: Bathysauroididae, Bathysauropsidae.
Comment: Suborders and families listed are as in Davis [7]. Although not monophyletic herein, the monophyly aulopiform suborders was supported by Davis [7].
Section Ctenosquamata (97%)
Subsection Myctophata (100%)
Order Myctophiformes
Myctophidae
Neoscopelidae
Subsection Acanthomorphata (97%)
Division Lampridacea (100%)
Order Lampridiformes, new circumscription
Lamprididae
Lophotidae
Regalecidae
Trachipteridae
Not examined: Radiicephalidae, Veliferidae.
Division Paracanthomorphacea sensu Grande et al. [20] (93%)
Series Percopsaria (100%)
Order Percopsiformes
Amblyopsidae
Aphredoderidae
Percopsidae
Series Zeiogadaria (=Zeioigadiformes sensu Li et al. [21]) (98%)
Subseries Zeariae (100%)
Order Zeiformes
Parazenidae
Zeidae
Not examined: Cyttidae, Grammicolepididae, Oreosomatidae, Zenionidae.
Subseries Gadariae (100%)
Order Stylephoriformes (sensu Miya et al. [22])
Stylephoridae
Order Gadiformes (100%)
Suborder Macrouroidei
Macrouridae (not monophyletic)
Steindachneriidae
Suborder Gadoidei (not monophyletic)
Gadidae
Lotidae
Merlucciidae
Moridae
Phycidae
Not examined: Bregmacerotidae, Euclichthyidae, Melanonidae.
Suborder Muraenolepidoidei
Not examined: Muraenolepididae.
Comment: The subordinal classification follows Roa-Varon and Orti [14: fig. 6].
Division Polymixiacea (100%)
Order Polymixiiformes
Polymixiidae
Division Euacanthomorphacea sensu [23] (99%)
Subdivision Berycimorphaceae (87%)
Order Beryciformes, new circumscription (circumscription similar to Trachichthyiformes sensu Moore [24])
Anoplogastridae
Barbourisiidae
Berycidae
Cetomimidae
Diretmidae
Melamphaidae
Monocentridae
Rondeletiidae
Trachichthyidae
Not examined: Anomalopidae, Gibberichthyidae, Hispidoberycidae, Stephanoberycidae (placement expected following Moore [24]).
Subdivision Holocentrimorphaceae (100%)
Order Holocentriformes, new
Holocentridae
Comment: Moore [24], and Stiassny and Moore [25] provide morphological evidence supporting a sister-group relationship between holocentrids and percomorphs, which further guarantees placement of this family in its own order.
Subdivision Percomorphaceae
Subdivision Percomorphaceae (cont.)
Not examined (9 families traditionally placed in “Perciformes”): Banjosidae, Bathyclupeidae, Dichistiidae, Hapalogenyidae, Lactariidae, Parascorpididae, Perciliidae, Symphysanodontidae, Trichonotidae.
Comment: These “perciform” families are provisionally placed here given the long history of phylogenetic indistinctiveness between Percoidei, Perciformes, and Percomorpha (e.g., [26]).
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 [27,28]) (100%)
Comment: In addition to the well-supported molecular circumscription of this group, kurtids, apogonids and gobioids are characterized by the presence of sensory papillae rows on the head and body [27].
Order Kurtiformes, new circumscription (98%)
Suborder Kurtoidei
Kurtidae
Suborder Apogonoidei
Apogonidae
Comment: Johnson [29] 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: Kraemeriidae, Rhyacichthyidae, Schindleriidae, Thalasseleotrididae, Xenisthmidae.
Series Scombrimorpharia (97%)
Order Syngnathiformes, new circumscription (97%)
Incertae sedis: Aulostomidae, Centriscidae.
Suborder Syngnathoidei, new circumscription (92%)
Syngnathidae
Possibly included (examined): Fistulariidae
Suborder Dactylopteroidei (100%)
Dactylopteridae
Suborder Callionymoidei (100%)
Callionymidae
Not examined: Draconettidae, Pegasidae, Solenostomidae, (assumed affinity with Callionymidae).
Suborder Mulloidei, new circumscription (92%)
Creediidae
Mullidae
Not examined: Leptoscopidae (assumed affinity with Creediidae [30]).
Order Scombriformes, new circumscription (=Stromateoidei sensu Li et al. [21]) (100%)
Ariommatidae
Bramidae
Caristiidae
Centrolophidae
Chiasmodontidae
Gempylidae
Icosteidae
Nomeidae
Pomatomidae
Scombridae (not monophyletic)
Scombrolabracidae
Stromateidae
Trichiuridae
“Stromateoid” families not examined: Amarsipidae, Scombropidae, Tetragonuridae; not examined but assumed affinity: Arripidae [31].
Comment: interfamilial resolution in Scombriformes is tenuous; circumscription of scombriform families into suborders (e.g., Scombroidei, Stromateoidei, Icostoidei) requires further work.
Series Carangimorpharia (94%)
Subseries Anabantomorphariae (=Anabantiformes sensu Li et al. [21]) (99%)
Order Synbranchiformes, new circumscription (98%)
Suborder Indostomoidei
Indostomidae
Suborder Synbranchoidei
Synbranchidae
Suborder Mastacembeloidei
Mastacembelidae
Not examined: Chaudhuriidae.
Order Anabantiformes (100%)
Suborder Anabantoidei (95%)
Anabantidae
Helostomatidae
Osphronemidae
Not examined: Badidae, Pristolepididae.
Suborder Channoidei (85%)
Channidae
Nandidae
Subseries Carangimorphariae (=Carangimorpha sensu Li et al. [21]) (100%)
Incertae sedis: Centropomidae (sensu Greenwood), Leptobramidae, Menidae, Polynemidae, Sphyraenidae, Toxotidae.
Order Istiophoriformes, new (100%)
Istiophoridae
Xiphiidae
Order Carangiformes sedis mutabilis (not monophyletic)
Carangidae
Coryphaenidae
Echeneidae
Nematistiidae
Rachycentridae
Comment: Monophyly of Carangiformes is not significantly rejected by the data (see also [4]).
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 recovered as the sister group of pleuronectoids in the present analysis, the order was resolved as monophyletic by a recent study, albeit with low support [4].
Subseries Ovalentariae (sensu Smith & Near in Wainwright et al. [3]; =Stiassnyiformes sensu Li et al. [21]) (100%)
Incertae sedis: Ambassidae, Embiotocidae, Grammatidae (in part; not monophyletic), Plesiopidae, Polycentridae, Pomacentridae, Pseudochromidae (not monophyletic).
Superorder Cichlomorphae (93%)
Order Cichliformes
Cichlidae
Order Pholidichthyiformes
Pholidichthyidae
Superoder Atherinomorphae (100%)
Order Atheriniformes (100%)
Atherinidae
Atherinopsidae
Bedotiidae
Isonidae
Melanotaeniidae
Pseudomugilidae
Telmatherinidae
Not examined: Dentatherinidae, Notocheiridae, Phallostethidae.
Order Beloniformes (32%)
Suborder Adrianichthyoidei
Adrianichthyidae
Suborder Exocoetoidei (100%)
Belonidae (not monophyletic)
Exocoetidae (not monophyletic)
Hemiramphidae (not monophyletic)
Scomberesocidae
Zenarchopteridae (not monophyletic)
Order Cyprinodontiformes (57%)
Suborder Aplocheiloidei
Aplocheilidae
Suborder Cyprinodontoidei (100%)
Cyprinodontidae
Fundulidae
Poeciliidae
Not examined: Anablepidae, Goodeidae, Nothobranchiidae, Profundulidae, Rivulidae, Valenciidae.
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. [21] plus Gobiesocidae) (100%)
Suborder Gobiesocoidei
Gobiesocidae
Subroder Blennioidei sedis mutabilis (not monophyletic)
Blenniidae
Chaenopsidae
Clinidae
Dactyloscopidae
Labrisomidae (not monophyletic)
Tripterygiidae
Comment: While blennioids are not monophyletic in Fig. S1, we note that preliminary analyses resulted in the reciprocal monophyly of gobiesocoids and blennioids, which is also congruent with a recent multi-locus analysis [3]. Monophyly of gobiesocoids and blennioids (as separate orders/suborders) is further supported by morphological evidence [1].
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.
Not examined: Datnioididae (assumed affinity with Lobotidae [11]); Callanthiidae, Cepolidae (see [21]); Dinolestidae, Dinopercidae, Ostracoberycidae, Pentacerotidae (see [26]).
Order Uranoscopiformes, new circumscription (=Paratrachiniformes sensu Li et al. [21]) (95%)
Ammodytidae
Pinguipedidae
Uranoscopidae
Not examined: Cheimarrichthyidae (see Li et al. [21]).
Order Labriformes sensu stricto (100%)
Labridae (not monophyletic)
Odacidae
Scaridae
Order Ephippiformes, new circumscription (100%)
Drepaneidae
Ephippidae
Comment: Greenwood et al. [32] hypothesized a close affinity between Drepane and ephippids.
Order Spariformes sensu Akazaki [33] and Johnson [34] (94%)
Lethrinidae
Sparidae
Possibly included (examined): Nemipteridae
Not examined but assumed affinity [34]: Centracanthidae
Comment: Akazaki [33] proposed that Lethrinidae
Sparidae, and Nemipteridae were closely related based on specializations of the suspensorium and other features [29]. Johnson [34] supported the monophyly of Akazaki's spariforms with the addition of Centracanthidae.
Order Lophiiformes (100%)
Suborder Lophioidei (100%)
Lophiidae
Suborder Antennarioidei (100%)
Antennariidae
Not examined: Brachionichthyidae, Lophichthyidae, Tetrabrachiidae.
Suborder Chaunacoidei (100%)
Chaunacidae
Suborder Ogcocephaloidei (100%)
Ogcocephalidae
Suborder Ceratioidei (not monophyletic)
Ceratiidae (not monophyletic)
Gigantactinidae
Himantolophidae
Melanocetidae
Oneirodidae
Not examined: Caulophrynidae, Centrophrynidae, Diceratiidae, Linophrynidae, Neoceratiidae, Thaumatichthyidae.
Comment: The monophyly of Ceratioidei is challenged in this analysis due to misplacement of Cryptoceratias; however, another recent mitogenomic study provided evidence for the monophyly of the suborder [35].
Order Tetraodontiformes (100%)
Suborder Triacanthodoidei (100%)
Triacanthodidae
Suborder Tetraodontoidei, new circumscription (100%)
Diodontidae
Tetraodontidae
Suborder Moloidei, new (100%)
Molidae
Suborder Balistoidei, new circumscription (100%)
Balistidae
Monacanthidae
Suborder Ostracioidei, new (100%)
Aracanidae
Ostraciidae
Suborder Triacanthoidei, new
Triacanthidae
Not examined or classified: Triodontidae.
Comment: Suborders recognized by Santini and Tyler [36] are not recovered here or in previous molecular studies and thus a new subordinal classification is proposed.
Order Acanthuriformes, restricted circumscription (see also [37]) (100%)
Acanthuridae
Luvaridae
Zanclidae
Order Pempheriformes, new (100%)
Glaucosomatidae
Pempheridae
Comment: Tominaga [38] suggested that features of the cranium and swimbladder may be homologous in Pempheris and Glaucosoma.
Order Cirrhitiformes, new circumscription (similar to Cirrhitoidea sensu Greenwood [39], and Burridge and Smolenski [40]) (91%)
Cheilodactylidae
Cirrhitidae
Not examined but expected affinity [39,40]: Aplodactylidae, Chironemidae, Latridae.
Order Centrarchiformes, new circumscription (100%)
Centrarchidae
Elassomatidae
Order Perciformes, new circumscription (=Serraniformes sensu Li et al. [21], Lautredou et al. [41]) (89%)
Incertae sedis: Platycephalidae, Percophidae.
Not examined (14 families traditionally placed in Scorpaeniformes): Apistidae, Aploactinidae, Bembridae, Champsodontidae, Congiopodidae, Cottocomephoridae, Eschmeyeridae, Gnathanacanthidae, Hoplichthyidae, Neosebastidae, Pataecidae, Perryenidae, Plectrogeniidae, Zanclorhynchidae.
Suborder Serranoidei sedis mutabilis (19%)
Serranidae
Suborder Percoidei, restricted circumscription (100%)
Percidae
Not examined: Trachinidae.
Comment: Lautredou et al. [41] 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)
Setarchidae
Synanceiidae
Tetrarogidae
Suborder Triglioidei sensu Jordan [42] (100%)
Triglidae
Peristediidae
Suborder Cottioidei, new circumscription (=Cottimorpha sensu Li et al. [21]) (100%)
Not examined: Trichodontidae (see [26])
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%)
Aulorhynchidae
Gasterosteidae
Hypoptychidae
Infraorder Zoarcales (100%)
Anarhichadidae
Bathymasteridae (not monophyletic)
Cryptacanthodidae
Stichaeidae (not monophyletic)
Pholidae
Zaproridae
Zoarcidae
Not examined: Ptilichthyidae, Scytalinidae.
Infraorder Cottales (96%)
Agonidae (not monophyletic)
Cyclopteridae
Cottidae (not monophyletic)
Hexagrammidae (not monophyletic)
Liparidae
Psychrolutidae
Not examined: 8 families traditionally placed in Cottoidei: Abyssocottidae, Bathylutichthyidae, Comephoridae, Ereuniidae, Hemitripteridae, Normanichthyidae, Parabembridae, Rhamphocottidae.
Superclass Sarcopterygii (96%)
Class Coelacanthimorpha (=Actinistia)
Order Coelacanthiformes
Latimeriidae
Class Dipnotetrapodomorpha sedis mutabilis (65%)
Subclass Dipnomorpha (100%)
Superorder Ceratodontae
Order Ceratodontiformes
Suborder Ceratodontoidei
Neoceratodontidae
Suborder Lepidosirenoidei (100%)
Lepidosirenidae
Protopteridae
Subclass Tetrapodomorpha (100%)
References
1. Wiley EO, Johnson GD (2010) A teleost classification based on monophyletic groups. In: Nelson JS, Schultze HP, Wilson MVH, editors. Origin and Phylogenetic Interrelationships of Teleosts. München, Germany: Verlag Dr. Friedrich Pfeil. pp. 123-182.
2. Arratia G (1999) The monophyly of Teleostei and stem-group teleosts. Consensus and disagreements. In: Arratia G, Schultze HP, editors. Mesozoic Fishes 2 – Systematics and Fossil Record. München: Verlag Dr. F. Pfeil. pp. 265-334.
3. Wainwright PC, Smith WL, Price SA, Tang KL, Sparks JS, et al. (2012) The evolution of pharyngognathy: a phylogenetic and functional appraisal of the pharyngeal jaw key innovation in labroid fishes and beyond. Systematic Biology 61: 1001-1027.
4. Betancur-R. R, Li C, Munroe TA, Ballesteros JA, Orti G (In press) Addressing gene-tree discordance and non-stationarity to resolve a multi-locus phylogeny of the flatfishes (Teleostei: Pleuronectiformes). Systematic Biology.
5. Eschmeyer WN, Fong JD (2013) Species of Fishes by family/subfamily. http://research.calacademy.org/ichthyology/catalog/family.
6. van der Laan R, Fricke R, Eschmeyer WN (2013) Family-Group Names. California Academy of Sciences http://research.calacademy.org/ichthyology/catalog/family.
7. Davis MP (2010) Evolutionary relationships of the Aulopiformes (Euteleostei: Cyclosquamata): a molecular and total evidence approach. In: Nelson JS, Schultze HP, Wilson MVH, editors. Origin and Phylogenetic Interrelationships of Teleosts. München, Germany: Verlag Dr. Friedrich Pfeil. pp. 317-336.
8. Greenwood PH (1976) A review of the family Centropomidae (Pisces, Perciformes). Bulletin of the British Museum (Natural History) Zoology 29: 1–81.
9. Li C, Betancur-R. R, Smith WL, Orti G (2011) Monophyly and interrelationships of Snook and Barramundi (Centropomidae sensu Greenwood) and five new markers for fish phylogenetics. Molecular Phylogenetics and Evolution 60: 463-471.
10. Chen WJ, Lheknim V, Mayden RL (2009) Molecular phylogeny of the Cobitoidea (Teleostei: Cypriniformes) revisited: position of enigmatic loach Ellopostoma resolved with six nuclear genes. Journal of Fish Biology 75: 2197-2208.
11. Nelson JS (2006) Fishes of the world. Hoboken: John Wiley & Sons. 601 p.
12. Sullivan JP, Lundberg JG, Hardman M (2006) A phylogenetic analysis of the major groups of catfishes (Teleostei : Siluriformes) using rag1 and rag2 nuclear gene sequences. Molecular Phylogenetics and Evolution 41: 636-662.
13. Li C, Orti G, Zhao J (2010) The phylogenetic placement of sinipercid fishes ("Perciformes") revealed by 11 nuclear loci. Molecular Phylogenetics and Evolution 56: 1096-1104.
14. Roa-Varon A, Orti G (2009) Phylogenetic relationships among families of Gadiformes (Teleostei, Paracanthopterygii) based on nuclear and mitochondrial data. Molecular Phylogenetics and Evolution 52: 688-704.
15. Chapleau F (1993) Pleuronectiform relationships - A cladistic reassessment. Bulletin of Marine Science 52: 516-540.
16. Munroe TA (2005) Chapter 3: Distributions and Biogeography. In: Gibson RN, editor. Flatfishes: Biology and Exploitation Blackwell Science Ltd. pp. 42-67.
17. Arratia G (2010) The Clupeocephala re-visited: Analysis of characters and homologies. Revista de Biologia Marina & Oceanografia 45: 635-657.
18. Albert JS, Crampton WGR (2005) Diversity and Phylogeny of Neotropical Electric Fishes (Gymnotiformes). In: Bullock CD, Hopkins AN, Fay RR, editors. Electroreception. New York: Springer. pp. 360–409.
19. Calcagnotto D, Schaefer SA, DeSalle R (2005) Relationships among characiform fishes inferred from analysis of nuclear and mitochondrial gene sequences. Molecular Phylogenetics and Evolution 36: 135-153.
20. Grande T, Borden WC, Smith WL (In press) Limits and relationships of Paracanthopterygii: A molecular framework for evaluating past morphological hypotheses. In: Arratia G, Schultze H-P, Wilson MVH, editors. Mesozoic Fishes 5 - Global Diversity and Evolution. Muenchen: Verlag F. Pfeil.
21. Li B, Dettai A, Cruaud C, Couloux A, Desoutter-Meniger M, et al. (2009) RNF213, a new nuclear marker for acanthomorph phylogeny. Molecular Phylogenetics and Evolution 50: 345-363.
22. Miya M, Holcroft NI, Satoh TP, Yamaguchi M, Nishida M, et al. (2007) Mitochondrial genome and a nuclear gene indicate a novel phylogenetic position of deep-sea tube-eye fish (Stylephoridae). Ichthyological Research 54.
23. Johnson GD, Patterson C (1993) Percomorph phylogeny: a survey of acanthomorphs and a new proposal. Bulletin of Marine Science 52: 554–626.
24. Moore JA (1993) Phylogeny of the Trachichthyiformes (Teleostei: Percomorpha). Bulletin of Marine Science 52: 114-136.
25. Stiassny MLJ, Moore JA (1992) A review of the pelvic girdle of atherinomorph fishes. Zoological Journal of the Linnean Society 104: 209-242.
26. Smith WL, Craig MT (2007) Casting the percomorph net widely: The importance of broad taxonomic sampling in the search for the placement of serranid and percid fishes. Copeia: 35-55.
27. Thacker C (2009) Phylogeny of Gobioidei and Placement within Acanthomorpha, with a New Classification and Investigation of Diversification and Character Evolution. Copeia 2009: 93-104.
28. Chakrabarty P, Davis MP, Sparks JS (2012) The First Record of a Trans-Oceanic Sister-Group Relationship between Obligate Vertebrate Troglobites. PLoS ONE.
29. Johnson GD (1993) Percomorph phylogeny: progress and problems. Bulletin of Marine Science 52: 3- 28.
30. Odani K, Imamura H (2011) New Phylogenetic Proposal for the Family Leptoscopidae (Perciformes: Trachinoidei). Bulletin of Fisheries Sciences Hokkaido University 61: 49-63.
31. Yagishita N, Miya M, Yamanoue Y, Shirai SM, Nakayama K, et al. (2009) Mitogenomic evaluation of the unique facial nerve pattern as a phylogenetic marker within the percifom fishes (Teleostei: Percomorpha). Molecular Phylogenetics and Evolution 53: 258-266.
32. Greenwood PH, Rosen DE, Weitzman SH, Myers GS (1966) Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bulletin of the American Museum of Natural History 131.
33. Akazaki M (1962) Studies on the spariform fishes-anatomy - phylogeny, ecology, and taxonomy. Osaka, Japan: Kosugi Co. Ltd. 368 p.
34. Johnson GD (1981) The limits and relationships of the Lutjanidae and associated families. Bulletin of the Scripps Institution of Oceanography 24: 1-114.
35. Miya M, Pietsch T, Orr J, Arnold R, Satoh T, et al. (2010) Evolutionary history of anglerfishes (Teleostei: Lophiiformes): a mitogenomic perspective. BMC Evolutionary Biology 10: 58.
36. Santini F, Tyler JC (2003) A phylogeny of the families of fossil and extant tetraodontiform fishes (Acanthomorpha, Tetraodontiformes), Upper Cretaceous to Recent. Zoological Journal of the Linnean Society 139: 565–617.
37. Holcroft NI, Wiley EO (2008) Acanthuroid relationships revisited: a new nuclear gene-based analysis that incorporates tetraodontiform representatives. Ichthyological Research 55: 274-283.
38. Tominaga Y (1986) The relationships of the families Glaucosomatidae and Pempherididae. In: Uyeno T, Arai R, Taniuchi T, Matsuura K, editors. Indo-Pacific fish biology. Tokyo: lchthyological Society of Japan. pp. 595-599.
39. Greenwood PH (1995) A revised familial classification for certain cirrhitoid genera (Teleostei, Percoidei, Cirrhitoidea), with comments on the groups monophyly and taxonomic ranking. Bulletin of the British Museum (Natural History) Zoology 61: 1–10.
40. Burridge CP, Smolenski AJ (2004) Molecular phylogeny of the Cheilodactylidae and Latridae (Perciformes: Cirrhitoidea) with notes on taxonomy and biogeography. Molecular Phylogenetics and Evolution 30: 118-127.
41. Lautredou A-C, Motomura H, Gallut C, Ozouf-Costaz C, Cruaud C, et al. (2013) Multi-scale exploration of the relationships among Serraniformes (Acanthomorpha, Teleostei) using new nuclear markers. Molecular Phylogenetics and Evolution 67: 140-155.
42. Jordan DS (1923) A classification of fishes including families and genera as far as known. Stanford Univ Publ, Univ Ser Biol Sci 3: 77-243.