Classification v.3







Phylogenetic Classification of Bony Fishes

Based mostly on Molecular Data — Version 3


 

Version Date: 31 July 2014 --- this version has been superseded by version 4 on 26 April 2017

Contributors to this version: Ricardo Betancur-R, Ed Wiley, Nicolas Bailly, Masaki Miya, Guillaume Lecointre, Guillermo Ortí


Citing this classification:

Betancur-R, R., E. Wiley, N. Bailly, M. Miya, G. Lecointre, and G. Ortí. 2014. Phylogenetic Classification of Bony Fishes --Version 3 (http://www.deepfin.org/Classification_v3.htm).

Betancur-R., R., R.E. Broughton, E.O. Wiley, K. Carpenter, J.A. Lopez, C. Li, N.I. Holcroft, D. Arcila, M. Sanciangco, J. Cureton, F. Zhang, T. Buser, M. Campbell, T. Rowley, J.A. Ballesteros, G. Lu, T. Grande, G. Arratia & G. Ortí. 2013. The tree of life and a new classification of bony fishes. PLoS Currents Tree of Life. 2013 Apr 18.


Download useful resources:

Go to the bottom of this page to download: Spreadsheet with complete classification scheme; molecular phylogeny used as basis for classification (from version 2, needs to be updated to reflect the current classification scheme); and figures with summary phylogenetic hypothesis displaying all major groups.


This classification is an update of version 1 (18 April 2013) and version 2 (27 Nov 2013), originally published by Betancur-R. et al. (2013a). This version fixes involuntary errors and omissions (thanks to numerous colleagues who provided feedback!) and incorporates new findings from the recent literature. It is based on the same phylogenetic results used for version 2, from analysis of a molecular data set with 1591 taxa (the data set combines evidence published by Betancur-R. et al. (2013a) and Near et al. (2013)). Families in the classification are linked to FishBase family pages (Froese and Pauly, 2013) except for cases where there are discrepancies in the recognition of valid families, noted and justified in each case in the classification.

Version 3 presents new names for series included in subdivision Percomorphaceae, to avoid confusion between this taxon and series Percomorpharia, and hence disambiguate the meaning of "percomorph". Percomorpharia is now referred to as series Eupercaria, and the other series names also have simpler endings (dropping the term "morph" but keeping a consistent ending with "aria"). Under this scheme, the word "percomorph" refers unambiguously to subdivision Percomorphaceae, retaining its original meaning as implied by Wiley and Johnson (2010), following Johnson and Patterson (1993). The common name "eupercarians" now is available for taxa in the clade containing Perciformes, Labriformes, Tetraodontiformes and others (i.e., the "new bush at the top" sensu Betancur-R. et al. (2013a)). Figure 1 below shows the new names.

A complete list of changes from version 2 can be found here.

General comments on the classification are here.

Figure 1. Phylogeny of the nine major percomorph clades included in Subdivision Percomorphaceae. Each major clade is recognized as a series.


Classification for Bony Fishes (version 3)

 

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 (1999) (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 (2010) 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 (1999) (38%)

Supercohort Osteoglossocephala sensu Arratia (1999) (99%)

Cohort Osteoglossomorpha

Order Hiodontiformes (100%)

Hiodontidae

Order Osteoglossiformes (100%)

Gymnarchidae

Mormyridae

Notopteridae

Osteoglossidae (includes Arapaima and Heterotis, formerly in Arapaimidae)

Pantodontidae

Supercohort Clupeocephala sensu Arratia (2010) (100%)

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

Subcohort Clupei (87%)

Order Clupeiformes

Suborder Denticipitoidei

Denticipitidae

Suborder Clupeoidei (100%)

Chirocentridae

Clupeidae (not monophyletic; includes Sundasalanx according to Lavoué et al. (2013), formerly in Sundalangidae)

Engraulidae

Pristigasteridae

Not examined: Dussumieriidae.

Comment: family-level groupings may require major revision; Pristigasteridae, Chirocentridae and Engraulidae are supported by other molecular studies, but not Clupeidae (Li and Ortí, 2007; Lavoué et al., 2013); five well-supported lineages identified by Lavoué et al. (2013) could become new families.

Subcohort Alepocephali (100%)

Order Alepocephaliformes

Alepocephalidae (not monophyletic)

Platytroctidae

Not examined: Leptochilichthyidae.

Comment: the position of alepocephaloids as the sister group to Ostariophyisi also was reported by Lavoué et al. (2008) and by Paulsen et al. (2009).

Subcohort Ostariophysi (100%)

Section Anotophysa (97%)

Order Gonorynchiformes

Suborder Gonorynchoidei

Gonorynchidae

Suborder Chanoidei

Chanidae

Suborder Knerioidei (100%)

Kneriidae (includes the former Phractolaemidae, now a subfamily in Kneriidae, following Davis et al. (2013))

Section Otophysa (100%)

Superorder Cypriniphysae (100%)

Order Cypriniformes

Botiidae (following Chen et al. (2009))

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 here, but see Alves-Gomes (2010) and Nakatani et al. (2011))

Suborder Sternopygoidei (not monophyletic)

Apteronotidae

Rhamphichthyidae

Sternopygidae (not monophyletic)

Not examined: Hypopomidae.

Comment: Although not monophyletic here, the monophyly of gymnotiform suborders is corroborated by Albert and Crampton (2005).

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 (Calcagnotto et al., 2005). In other molecular studies, the monophyly of Characiformes was not obtained, since Characoidei was more closely related to Siluriformes than to Citharinoidei (Nakatani et al., 2011; Chen et al., 2014a).

Order Siluriformes (100%)

Suborder Loricarioidei (93%)

Astroblepidae

Callichthyidae

Loricariidae

Nematogenyidae

Trichomycteridae

Not examined: Scoloplacidae

Suborder Diplomystoidei

Diplomystidae

Suborder Siluroidei (100%)

Akysidae

Amblycipitidae

Amphiliidae

Anchariidae

Ariidae

Auchenipteridae

Bagridae (includes taxa formerly in Olyridae, following Sullivan et al. (2006))

Cetopsidae

Chacidae

Clariidae

Claroteidae

Cranoglanididae

Doradidae

Heptapteridae

Heteropneustidae

Horabagridae (following Sullivan et al. (2006))

Ictaluridae

Malapteruridae

Mochokidae

Pangasiidae

Pimelodidae

Plotosidae

Pseudopimelodidae

Schilbeidae

Siluridae

Sisoridae

Not examined: Aspredinidae, Austroglanididae, Erethistidae, Lacantuniidae. 

Supercohort Clupeocephala (cont.)

Cohort Euteleosteomorpha (100%)

Subcohort Lepidogalaxii

Order Lepidogalaxiiformes

Lepidogalaxiidae

Subcohort Protacanthopterygii sedis mutabilis (37%)

Order Galaxiiformes (100%)

Galaxiidae

Order Argentiniformes (100%)

Argentinidae

Bathylagidae

Microstomatidae

Opisthoproctidae

Order Salmoniformes (100%)

Salmonidae

Order Esociformes (100%)

Esocidae

Umbridae

Subcohort Stomiati (73%)

Order Stomiatiformes (=Stomiiformes) (100%)

Diplophidae (following Nelson (2006); apparently ommited or considerd a junior synonym of Gonostomidae by Eschmeyer and Fong (2014))

Gonostomatidae

Phosichthyidae (not monophyletic)

Sternoptychidae

Stomiidae

Order Osmeriformes (100%)

Osmeridae

Plecoglossidae

Retropinnidae

Salangidae

Subcohort Neoteleostei (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 (includes taxa previously listed in Omosudidae and Anotopteridae, following Davis (2010))

Bathysauridae

Chlorophthalmidae (not monophyletic)

Evermannellidae

Giganturidae

Ipnopidae (not monophyletic)

Notosudidae

Paralepididae (not monophyletic)

Scopelarchidae (not monophyletic)

Sudidae (following Davis (2010); synonym of Paralepididae according to Eschmeyer (2014))

Not examined: Bathysauroididae, Bathysauropsidae sensu Davis (2010).

Comment: Anotopteridae is not recognized; aulopiform families listed follow Davis (2010). Although not monophyletic herein, the monophyly of aulopiform suborders listed is supported by Davis (2010).

Section Ctenosquamata (97%)

Subsection Myctophata (100%)

Order Myctophiformes

Myctophidae

Neoscopelidae

Subsection Acanthomorphata (97%)

Division Lampridacea (100%)

Order Lampridiformes

Lamprididae

Lophotidae

Regalecidae

Trachipteridae

Not examined: Radiicephalidae, Veliferidae.

Division Paracanthomorphacea sensu Grande et al. (2013) (93%)

Series Percopsaria (100%)

Order Percopsiformes

Amblyopsidae

Aphredoderidae

Percopsidae

Series Zeiogadaria (= Zeiogadiformes sensu Li et al. (2009)) (98%)

Subseries Zeariae (100%)

Order Zeiformes

Parazenidae

Zeidae

Zenionidae

Not examined: Cyttidae, Grammicolepididae, Oreosomatidae.

Subseries Gadariae (100%)

Order Stylephoriformes (sensu Miya et al. (2007))

Stylephoridae

Order Gadiformes (100%)

Suborder Macrouroidei

Macrouridae (not monophyletic here, but see Roa-Varon and Ortí (2009))

Steindachneriidae (following Roa-Varon and Ortí (2009); synonym of Merluciidae according to Eschmeyer (2014))

Suborder Gadoidei (not monophyletic)

Gadidae

Gaidropsaridae (formerly a subfamily of Lotidae, raised to family level)

Lotidae

Merlucciidae

Moridae

Phycidae

Not examined: Bregmacerotidae, Euclichthyidae, Melanonidae.

Suborder Muraenolepidoidei

Muraenolepididae

Comment: The subordinal classification follows Roa-Varón and Ortí (2009): fig. 6.

Division Polymixiacea (100%)

Order Polymixiiformes

Polymixiidae

Division Euacanthomorphacea sensu Johnson and Patterson (1993) (99%)

Subdivision Berycimorphaceae (87%)

Order Beryciformes (similar to Trachichthyiformes sensu Moore (1993))

Anomalopidae

Anoplogastridae

Barbourisiidae

Berycidae

Cetomimidae

Diretmidae

Melamphaidae

Monocentridae

Rondeletiidae

Stephanoberycidae

Trachichthyidae (not monophyletic)

Not examined: Gibberichthyidae, Hispidoberycidae (expected affinity following Moore (1993)).

Subdivision Holocentrimorphaceae (100%)

Order Holocentriformes

Holocentridae

Comment: Moore (1993) and Stiassny and Moore (1992) 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 (="Percomorpha" sensu Miya et al. (2003) and Miya et al. (2005)).

Subdivision Percomorphaceae (cont.)

Series Ophidiaria (100%)

Order Ophidiiformes

Suborder Ophidioidei

Carapidae

Ophidiidae (not monophyletic as traditionally recognized, but monophyletic if Carapidae is included)

Suborder Bythitoidei

Aphyonidae

Bythitidae (not monophyletic)

Not examined: Parabrotulidae

Series Batrachoidaria (100%)

Order Batrachoidiformes

Batrachoididae

Series Gobiaria (see also Thacker (2009) and Chakrabarty et al. (2012)) (100%)

Order Kurtiformes (98%)

Suborder Kurtoidei

Kurtidae

Suborder Apogonoidei

Apogonidae

Comment: Johnson (1993) noted that the configuration of 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 (includes Microdesmus, previously listed in Microdesmidae; see Thacker (2009))

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

Comment: In addition to the well-supported molecular circumscription of Gobiaria, kurtids, apogonids and gobioids are characterized by the presence of sensory papillae rows on the head and body (Thacker, 2009).

Series Syngnatharia (96%)         

Order Syngnathiformes

Suborder-level incertae sedis in Syngnathiformes

Aulostomidae

Centriscidae

Fistulariidae

Suborder Syngnathoidei (92%)

Syngnathidae

Pegasidae

Not examined: Solenostomidae (assumed affinity with Syngnathidae)

Suborder Dactylopteroidei (100%)

Dactylopteridae

Suborder Callionymoidei (100%)

Callionymidae

Not examined: Draconettidae (assumed affinity with Callionymidae).

Suborder Mulloidei (92%)

Mullidae

Series Pelagiaria (= Stromateoidei sensu Li et al. (2009); = Pelagia sensu Miya et al. (2013)) (100%)          

Order Scombriformes

Ariommatidae

Arripidae (not examined, included here following Yagishita et al. (2009) and Miya et al. (2013))

Bramidae

Caristiidae

Centrolophidae

Chiasmodontidae

Gempylidae (not monophyletic)

Icosteidae

Nomeidae

Pomatomidae

Scombridae (not monophyletic here, but see Miya et al. (2013))

Scombrolabracidae

Stromateidae

Trichiuridae

Families not examined: Amarsipidae, Scombropidae (Doiuchi et al. (2004); Miya et al. (2013)), Tetragonuridae (not examined, included here following Miya et al. (2013))

Comment: interfamilial resolution in Scombriformes is tenuous; circumscription of scombriform families into suborders (e.g., Scombroidei, Stromateoidei, Icostoidei) or new orders requires further work.

Series Anabantaria (= Anabantiformes sensu Li et al. (2009)) (99%)

Order Synbranchiformes (100%)

Suborder Mastacembeloidei

Mastacembelidae

Not examined: Chaudhuriidae.

Suborder Indostomoidei

            Indostomidae

Suborder Synbranchoidei

Synbranchidae

Order Anabantiformes (= Labyrinthici) (100%)

Suborder Anabantoidei (95%), new circumscription

Anabantidae

Helostomatidae

Osphronemidae

Suborder Channoidei (100%), new circumscription

Channidae

Suborder Nandoidei (94%), new anabantiform suborder

Badidae

Nandidae

Pristolepididae

Comment: suborders of Anabantiformes now reflect well-supported monophyletic groups, correcting an error in Version 2 and defining a third suborder (Channoidei) for the family Channidae. Affinities of Channidae with other anabantiform families varies among studies (e.g., Near et al. (2013), Betancur-R. et al. (2013a), and our new results). The new scheme with three subroders is robust to this ambiguity.

Series Carangaria (= Carangimorpha sensu Li et al. (2009)) (100%)

Order-level incertae sedis in Carangiaria

Centropomidae (includes taxa formelry listed in Latidae, following Greenwood (1976) and Li et al. (2011))

Leptobramidae

Menidae

Polynemidae

Sphyraenidae

Toxotidae

Not examined: Lactariidae (included in Carangimorphariae according to Campbell et al. (2013))

Order Istiophoriformes (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 (Betancur-R. et al., 2013b).

Order Pleuronectiformes sedis mutabilis (not monophyletic)

Suborder Psettodoidei (100%)

Psettodidae

Suborder Pleuronectoidei (100%)

Achiridae

Bothidae

Citharidae

Cynoglossidae

Paralichthyidae (not monophyletic as traditionally recognized, but monophyletic if the Cyclopsetta group is included in its own family; see Betancur-R. et al. (2013b))

Pleuronectidae

Poecilopsettidae (following Chapleau (1993) and Munroe (2005))

Rhombosoleidae (following Chapleau (1993) and Munroe (2005); includes taxa listed in Achiropsettidae according to Betancur-R. et al. (2013b))

Samaridae

Scophthalmidae

Soleidae

Not examined: Paralichthodidae (following Chapleau (1993) and Munroe (2005)).

Comment: Although Psettodidae is not recovered as the sister group of pleuronectoids in the present analysis, the order was resolved as monophyletic by recent studies that address this issue specifically (Betancur-R. et al. (2013b), Betancur-R and Ortí (2014)).

Series Ovalentaria (sensu Smith and Near in Wainwright et al. (2012); = Stiassnyiformes sensu Li et al. (2009)) (100%)

Order-level incertae sedis in Ovalentaria

Ambassidae

Congrogadidae sensu Godkin and Winterbottom (1985) and Wainwright et al. (2012), formerly a subfamily of Pseudochromidae

Embiotocidae

Grammatidae (not monophyletic)

Opistognathidae

Plesiopidae

Polycentridae

Pomacentridae

Pseudochromidae

Superorder Cichlomorphae (93%)

Order Cichliformes

Cichlidae

Order Pholidichthyiformes

Pholidichthyidae

Superoder Atherinomorphae (100%)

Order Atheriniformes (100%)

Atherinidae

Atherinopsidae

Bedotiidae

Isonidae

Melanotaeniidae

Phallostethidae

Pseudomugilidae

Telmatherinidae

Not examined: Dentatherinidae, Notocheiridae.

Order Beloniformes (32%)

Suborder Adrianichthyoidei

Adrianichthyidae

Suborder Belonoidei (100%) (corrected name, listed as Exocoetoidei in version 2)

Belonidae (including the former Scomberesocidae; see Lovejoy and Collette (2001) and Lovejoy et al. (2004))

Exocoetidae

Hemiramphidae (not monophyletic)

Zenarchopteridae (not monophyletic here, but see Lovejoy et al. (2004))

Order Cyprinodontiformes (57%)

Suborder Aplocheiloidei

Aplocheilidae

Not examined: Nothobranchiidae, Rivulidae.

Suborder Cyprinodontoidei (100%)

Cyprinodontidae

Fundulidae

Poeciliidae

Not examined: Anablepidae, Goodeidae, Profundulidae, Valenciidae.

Superorder Mugilomorphae (100%)

Order Mugiliformes

Mugilidae

Superorder Blenniimorphae (80%)

Order Blenniiformes (100%) sensu Li et al. (2009)

Suborder Gobiesocoidei

Gobiesocidae

Suborder Blennioidei (not monophyletic; = Blenniiformes sensu Lin and Hastings (2013))

Blenniidae

Chaenopsidae

Clinidae

Dactyloscopidae

Labrisomidae (includes Stathmonotus; not monophyletic here, but see Lin and Hastings (2013))

Tripterygiidae

Comment: While blennioids are not monophyletic in our results, we note that preliminary analyses resulted in the reciprocal monophyly of gobiesocoids and blennioids, which is congruent with molecular (Wainwright et al., 2012; Lin and Hastings, 2013) and morphological (Springer and Orrell, 2004) evidence. Monophyly of gobiesocoids and blennioids (as separate orders/suborders) is further supported by both morphological (Wiley and Johnson, 2010) and molecular evidence. Chaenopsidae is monophyletic if Stathmonotus is included in Labrisomidae, following Lin and Hastings (2013).

Series Eupercaria (99%)

Order-level incertae sedis in Eupercaria

Callanthiidae

Caproidae

Champsodontidae

Cepolidae

Centrogenyidae

Emmelichthyidae

Gerreidae

Haemulidae

Lutjanidae (includes the former Caesionidae, following Johnson (1993), Miller and Cribb (2007), and others)

Malacanthidae

Monodactylidae

Moronidae

Pomacanthidae

Priacanthidae

Scatophagidae

Sciaenidae

Siganidae

Sillaginidae

Not examined: Dinolestidae, Dinopercidae (see Smith and Craig (2007)). Six families traditionally placed in “Perciformes” are also provisionally listed here are: Bathyclupeidae, Dichistiidae, Hapalogenyidae, Parascorpididae, Symphysanodontidae, Trichonotidae; these are not placed in the recently circumscribed Perciformes given the long history of phylogenetic indistinctiveness between Percoidei, Perciformes, and Percomorpha (e.g., Smith and Craig (2007)).

Order Uranoscopiformes (= Paratrachinoidei sensu Li et al. (2009)) (95%)

Ammodytidae

Cheimarrichthyidae

Pinguipedidae

Uranoscopidae

Possibly included: Centrogenyidae (bootstrap support for Uranoscopiformes plus Centrogenyidae is only 41%, as opposed to 99% for Uranoscopiformes s.s. but placement is congruent with results of Near et al. (2013).

Order Labriformes sensu stricto (100%)

Labridae (includes taxa previoulsy listed in Scaridae; see Price et al. (2010, 2011) and Wainwright et al. (2012))

Odacidae

Order Lobotiformes (100%)

Datnioididae

Lobotidae

Order Ephippiformes (100%)

Drepaneidae

Ephippidae

Comment: Greenwood et al. (1966) hypothesized a close affinity between Drepane and ephippids.

Order Spariformes sensu Akazaki (1962) and Johnson (1981) (94%)

Lethrinidae

Sparidae

Possibly included (examined): Nemipteridae

Not examined but assumed affinity (Johnson, 1981): Centracanthidae

Comment: Akazaki (1962) proposed that Lethrinidae, Sparidae, and Nemipteridae were closely related based on specializations of the suspensorium and other features (Johnson (1993)). Johnson (1981) supported the monophyly of Akazaki's spariforms with the addition of Centracanthidae.

Order Chaetodontiformes, new eupercarian order (50%)

Chaetodontidae

Leiognathidae

Comment: Although support for this clade is weak (50%) in our analysis, it has been consistently obtained by previous studies with higher nodal support (90-99% in Near et al. (2012a) and 70-89% in Near et al. (2013)).

Order Lophiiformes (100%). This order is the sister group of Tetraodontiformes (55% bootstrap); also supported by anatomical evidence (Chanet et al., 2013), larval characters (Baldwin, 2013), and previous molecular studies (e.g. Dettaï and Lecointre, 2008; Miya et al., 2003; Miya et al., 2010).

Suborder Lophioidei (100%)

Lophiidae

Suborder Antennarioidei (100%).

Antennariidae

Not examined: Brachionichthyidae, Lophichthyidae, Tetrabrachiidae.

Suborder Chaunacoidei (100%)

Chaunacidae

Suborder Ogcocephaloidei (100%)

Ogcocephalidae

Suborder Ceratioidei

Ceratiidae

Gigantactinidae

Himantolophidae

Melanocetidae

Oneirodidae

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

Order Tetraodontiformes (100%). This order is the sister group of Lophiiformes (55% bootstrap); also supported by anatomical evidence (Chanet et al., 2013), larval characters (Baldwin, 2013), and previous molecular studies (e.g. Dettaï and Lecointre 2008; Miya et al., 2003; Miya et al., 2010).

Suborder Triacanthodoidei (100%)

Triacanthodidae

Suborder Triacanthoidei

Triacanthidae

Suborder Tetraodontoidei (100%)

Diodontidae

Tetraodontidae

Suborder Moloidei (100%)

Molidae

Suborder Balistoidei (100%)

Balistidae

Monacanthidae

Suborder Ostracioidei (100%)

Aracanidae

Ostraciidae

Suborder Triodontoidei

Triodontidae

Comment: This subordinal classification differs from that proposed by Santini and Tyler (2003).

Order Acanthuriformes, restricted circumscription (see also Holcroft and Wiley (2008)) (100%)

Acanthuridae

Luvaridae

Zanclidae

Order Pempheriformes sedis mutabilis (44%)

Acropomatidae (not monophyletic)

Banjosidae

Creediidae

Epigonidae

Glaucosomatidae

Howellidae

Lateolabracidae

Ostracoberycidae

Pempheridae

Pentacerotidae

Percophidae

Polyprionidae (not monophyletic)

Not examined: Leptoscopidae (assumed affinity with Creediidae according to Odani and Imamura (2011))

Comment: Tominaga (1986) suggested that features of the cranium and swimbladder may be homologous in Pempheris and Glaucosoma. Although support for Pempheriformes is only 44%, this clade is often recovered in different analyses.

Order Centrarchiformes (77%), new circumscription

Suborder Centrarchoidei (89%)

Centrarchidae

Elassomatidae

Enoplosidae

Sinipercidae (following Li et al. (2010); synonym of Percichthyidae according to Eschmeyer (2014))

Comment: inclusion of Enoploside in this suboorder differs from results obtained by Lavoué et al. (2014).

Suborder Cirrhitioidei (similar to Cirrhithoidea sensu Greenwood (1995) and Burridge and Smolenski (2004); treated as Cirrhitiformes in previous versions of the classification) (81%)

Cheilodactylidae

Cirrhitidae

Not examined but expected affinity (Burridge and Smolenski, 2004; Greenwood, 1995): Aplodactylidae, Chironemidae, Latridae.

Suborder Percichthyoidei

Percichthyidae (not monophyletic, a new family for Percalates is necessary; includes Percilia)

Comment: percichthyoids and Percichthyidae sensu Johnson (1984) are not monophyletic: the Australian species Percalates colonorum and Percalates novemaculeata are not closely related to other members of Percichthyidae (Betancur-R. et al. (2013a); Chen et al. (2014b); Lavoué et al. (2014)), so these species are herein placed in their own suborder (Peter Unmack pers. comm.; Lavoué et al. (2014)). Percalates is listed as a junior synonym of Macquaria by Eschmeyer (2014), but the type species of Macquaria (M. australasica) is closely related to other species of Macquaria (M. ambigua) within Percichthyidae sensu stricto, hence both names are valid genera (Peter Unmack et al., pers. comm.; Lavoué et al. (2014)). Percichthyidae sensu stricto includes Percilia (formerly placed in its own family Perciliidae).

Suborder Percalatoidei, new

"Percalatidae" (to be described)

Comment: Formal description of a new family for Percalates is required to comply with the ICZN.

Suborder Terapontoidei (= Clade "h2" of Yagishita et al. (2009); treated as Terapontiformes in previous versions of the classification) (95%)

Girellidae (following Carpenter 2001; listed as a subfamily of Kiphosidae by Eschmeyer (2014))

Kuhliidae

Kyphosidae

Oplegnathidae

Terapontidae

Comment: Although the family name Cirrithidae Macleay 1841 is older than Centrarchidae Bleeker 1859, we retain the name Centrarchiformes for this order in agreement with previous usage but expand its membership following recent proposals by Near et al. (2013) and Chen et al. (2014b) and Lavoué et al. (2014).

Order Perciformes (= Serraniformes sensu Lautredou et al. (2013); Li et al. (2009) (99%)

Not examined (10 families traditionally placed in Scorpaeniformes): Apistidae, Aploactinidae, Congiopodidae, Eschmeyeridae, Gnathanacanthidae, Neosebastidae, Pataecidae, Perryenidae, Plectrogeniidae, Zanclorhynchidae.

Suborder-level incertae sedis in Perciformes

Bembropidae (following Smith and Craig (2007); synonym of Percophidae according to Eschmeyer (2014))

Hoplichthyidae

Platycephalidae.

Suborder Serranoidei sedis mutabilis (49%)

            Serranidae

Suborder Percoidei, restricted circumscription (99%)

            Niphonidae (following Smith and Craig (2007); synonym of Serranidae according to Eschmeyer (2014))

Percidae

            Not examined: Trachinidae.

Comment: Lautredou et al. (2013) using seven nuclear markers obtained a clade uniting Percidae and Trachinidae with full support.

Suborder Notothenioidei (100%)

Artedidraconidae

Bathydraconidae (not monophyletic)

Bovichtidae

Channichthyidae

Eleginopsidae

Harpagiferidae

Nototheniidae (not monophyletic)

Pseudaphritidae

Suborder Scorpaenoidei (83%)

            Scorpaenidae (not monophyletic)

            Sebastidae

            Setarchidae

Synanceiidae

Tetrarogidae

Suborder Bembroidei (96%)

Bembridae

Parabembridae

Suborder Triglioidei sensu Jordan (1923) (100%)

Peristediidae

Triglidae

Suborder Cottoidei (= Cottimorpha sensu Li et al. (2009)) (100%)

Comment: We have chosen to recognize clades within this suborder as infraorders, adopting the ending "–ales" for this rank. Gasterosteales and Zoarcales are probably sister-groups (although not in our results); they have been grouped as Zoarciformes by Li et al. (2009).

Infraorder Anoplopomatales (= Anoplopomatoidei in previous classifications)

Anoplopomatidae

Infraorder Gasterosteales (= Gasterosteoidei in previous classifications--in part, excluding Indostomidae) (100%)

Aulorhynchidae

Gasterosteidae

Hypoptychidae

Infraorder Zoarcales (= Zoarcoidei in previous classifications) (100%)

Anarhichadidae

Bathymasteridae (not monophyletic)

Cryptacanthodidae

Stichaeidae (not monophyletic)

Pholidae

Zaproridae

Zoarcidae

Not examined: Ptilichthyidae, Scytalinidae.

Infraorder Hexagrammales (100%) (= Hexagrammoidei in previous classifications)

Hexagrammidae (sensu stricto, following Smith and Busby (2014))

Comment: Hexagrammidae as formerly defined is not monophyletic. We now split it into two families (formerly subfamilies): Hexagrammidae (sensu stricto) and Zaniolepidoales following Washington et al. (1984), Shinohara (1994), and Smith and Busby (2014). As in previous cottoid classifications, these families are placed in their own infraorders (note that previous classifications use suborders instead of infraorders).

Infraorder Zaniolepidoales (= Zaniolepidoidei sensu Smith and Busby (2014))

Zaniolepididae (sensu Smith and Busby (2014), formerly a subfamily in Hexagrammidae; not listed by Eschmeyer (2014))

Infraorder Cottales (96%) (= Cottoidei sensu Smith and Busby (2014))

Agonidae (not monophyletic here but see Smith and Busby (2014); includes the former Hemitripteridae)

Cottidae (not monophyletic here but see Smith and Busby (2014); includes the former Abyssocottidae, Comephoridae, and Cottocomephoridae.)

Cyclopteridae

Liparidae

Psychrolutidae (includes the former Bathylutichthyidae)

Scorpaenichthyidae (see Smith and Busby (2014))

Trichodontidae

Not examined: Jordaniidae (not listed by Eschmeyer (2014); newly defined by Smith and Busby (2014)), Ereuniidae, Normanichthyidae, and Rhamphocottidae.

Comment: Smith and Busby (2014) changed the membership of Cottidae and Psycholutridae to achieve reciprocal monopyhyly of these families. Our phylogenetic results do not resolve a monophyletic Cottidae even under the new circumscription proposed by these authors. We refer the readers to that study for more details.

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%)

 


General comments on the classification

This classification (version 3), as its previous versions, also buids on Wiley and Johnson (2010) and Betancur-R. et al. (2013a), intending to preserve names and taxonomic composition of groups as much as possible. However, adjustments have been made to recognize well-supported molecular clades, many of which also have been obtained by previous molecular studies (several examples discussed below--references are cited when decisions are based on other publications). Order-level or supraordinal taxa have been erected (listed as new) or resurrected on the basis of well-supported clades only (>90% bootstrap values). Current taxon names supported by previous molecular or morphological studies have been retained if congruent with our results, even if bootstrap support is low (e.g., Osteoglossocephalai sensu Arratia (1999) 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., 2012) and the order Pleuronectiformes (not monophyletic here but monophyletic in Betancur-R. et al., 2013b). The classification is presented in phylogenetic order up to the subordinal rank (following the branching order in our results), but families within orders (or suborders) are listed alphabetically.

A total of 67 orders are classified, of which one is new (Chaetodontiformes) and two were sinked (Terapontiformes and Cirrhitiformes) in version 3. Three orders were new in the version 1 (Holocentriformes, Istiophoriformes, and Pempheriformes) and two were new in version 2 (Lobotiformes and Terapontiformes). The ordinal status of several percomorph families examined (as well as many others unexamined) belonging to the Series Carangimorpharia, Ovalentaria, and Percomorpharia remains uncertain (incertae sedis) due to poor phylogenetic resolution. We therefore list these families as incertae sedis within each of these groups (Carangimorpharia, Ovalentaria, and Percomorpharia) awaiting new phylogenetic evidence to clarify their ordinal status.

Family names for bony fishes are based on Eschmeyer and Fong (2014) and van der Laan et al. (2013), with minor modifications, indicted in each case in the classification. Van der Laan et al. (2013) and Eschmeyer (2014) should be consulted for authorship of family names and Wiley and Johnson (2010) 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 lists based on published studies that we know validate or synonymize taxa using explicit phylogenetic evidence. In order to minimize the number of non-monophyletic taxa, we have changed the membership of many families traditionally recognized in ichthyology whose validity is strongly challenged by molecular/morphological phylogenetic analyses. For instance, we no longer recognize families such as Scaridae, Caesionidae, and Microdesmidae (lumped with Labridae, Lutjanidae, and Gobiidae, respectively). Accordingly, the number of non-monophyletic families decreased from 41 in version 2 to 30 in this version (indicated in each case).

A total of 496 families of bony fishes are now recognized (excluding tetrapods), of which 394 (79.4%) were examined. 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 102 unexamined families can be easily obtained from this spreadsheet that also contains the complete classification, and is intended as a resource to help fish systematists direct future sequencing efforts.

 

The new classification scheme presented here (version 3) should be considered work in progress 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 becomes available and feedback from experts help refine it. Please send comments or concerns to classification@deepfin.org. For the most updated version always visit DeepFin.

 


List of all changes from version 2 to version 3

  • Name changes for Series in Percomorphaceae: new names are Ophidiaria, Batrachoidaria, Gobiaria, Syngnatharia, Pelagiaria, Anabantaria, Carangaria, Ovalentaria, and Eupercaria.
  • Definition of Centrarchiformes now follows Near et al. (2013), Lavoué et al. (2014), and Chen et al. (2014b) to include the families Centrarchidae, Elassomatidae, Sinipercidae Jordan, Percichthyidae, Enoplosidae, Cirrithidae, Cheilodactylidae, Girellidae, Kuhliidae, Kyphosidae, Oplegnathidae, and Terapontidae. This change sinks the orders Terapontiformes and Cirrhitiformes (recognized in previous versions), since they are now recognized as suborders of Centrarichiformes together with Centrarchoidei and Percichthyoidei. The new ordinal circumscription is relatively well supported (bootstrap value of 77%). Since Percilia is deeply nested within Percichthyidae in our analyses as well as in other studies mentioned above, the family Percilidae is no longer recognized as valid.
  • Suborders of Anabantiformes now reflect well-supported monophyletic groups, correcting an error in Version 2 and defining a third suborder (Channoidei) for the family Channidae. Affinities of Channidae with other anabantiform families varies among studies (e.g., Near et al. (2013), Betancur-R. et al. (2013a), and our new results). The new scheme with three subroders is robust to this ambiguity.
  • Family Arapaimidae no longer recognized (now included in Osteoglossidae).
  • Suborders in Beloniformes: Belonoidei replaces Exocoetoidei.
  • Notobranchidae, Rivulidae, and Aplocheilidae are removed from “not examined” in Cyprinodontoidei and listed as “not examined” in Aplocheiloidei.
  • Scoloplacidae is removed from “not examined” in Siluroidei and listed as “not examined” in Loricaroidei.
  • Family Scaridae is now included in Labridae, rendering Labridae monophyletic (citations???).
  • Centrogenyidae is removed from order-level incertae sedis in Eupercaria and listed as “possibly included” in Uranoscopiformes. Bootstrap support for Uranoscopiformes plus Centrogenyidae is only 41% (as opposed to 99% for Uranoscopiformes s.s.) but placement is congruent with results reported by Near et al. (2013).
  • Cottoidei was recently revised by Smith and Busby (2014) who proposed a revised classification for all families within this suborder except the Aulorhynchidae, Gasterosteidae, and Hypotychidae (infraorder Gasterosteales), that were not examined. We follow these authors to make the following changes in the classification: (1) Hexagrammidae is split it into two families, defined as subfamilies by previous authors (e.g., Washington et al. 1984; Shinohara 1994): Hexagrammidae (sensu stricto) and Zaniolepididae. Both families are now placed in their own infraorders (Zaniolepidoales and Hexagrammales), following previous cottoid classifications. Note that these classifications use suborder instead of infraorders as presented here. (2) We now recognize the cottoid families Jordaniidae (not examined) and Scorpaenichthyidae.

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