Classification v.4

Phylogenetic Classification of Bony Fishes

Version 4 


Version Date: XXX 2016

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

Cite this classification:

Betancur-R, R., E. Wiley, N. Bailly, A. Acero, M. Miya, G. Lecointre, and G. Ortí. 2016. Phylogenetic Classification of Bony Fishes – Version 4.

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

  • Annotated spreadsheet with the complete classification scheme (XLSX format
  • Phylogenetic tree used as the basis for this classification (newick format; and annotated PDF format)
  • Summary phylogeny displaying all major groups (PDF format; it can be edited in Adobe Illustrator or similar)

General comments on the classification

This classification is an update of previous versions (version 1version 2 and version 3)originally published by Betancur-R. et al. (2013a) and based on Wiley and Johnson (2010). This version incorporates new findings from recent studies and fixes involuntary errors and omissions. The phylogenetic framework is based on a recent update of the fish tree (Betancur-R. et al., 2015) with two clades grafted into it from two large-scale phylogenetic studies on percomorphs (Sanciangco et al., 2016) and otophysans (Arcila et al., 2017). The complete tree includes 1770 species of bony fishes and two chondrichthyian outgroups. 

This classification intends 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 recent studies. A total of 72 orders and 84 suborders of bony fishes are classified in this version (vs. 66 orders in version 1). 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 external phylogenetic evidence. Order-level or supraordinal taxa have been resurrected on the basis of well-supported clades. Current taxon names supported by previous molecular or morphological studies have been retained if congruent with our results, even if support values are low. In some cases, order-level taxa that were not monophyletic in our tree are also validated, as long as the incongruence is not substantially rejected by the data (e.g., poorly supported incongruent clades). The classification is presented in phylogenetic order up to the subordinal rank (following the branching order in our tree), but families within orders and suborders are listed alphabetically.

A total of 499 families of bony fishes are now recognized (excluding tetrapods), of which 397 (80%) were included in our large-scale phylogeny. Family names for bony fishes are based on and van der Laan et al. (2014) and Eschmeyer and Fong (2016), with modifications indicated in each case in the classification and on Table 2Van der Laan et al. (2014) should be consulted for authorship of family names. 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. Families in the classification are linked to FishBase family pages (Froese and Pauly, 2015) except for cases where discrepancies in the recognition of valid families exist, noted and justified in each case. 

In order to minimize the number of non-monophyletic taxa, we have changed the membership of some traditionally recognized families whose validity is strongly challenged by phylogenetic evidence. For instance, we no longer recognize families such as Carapidae, Scaridae, Caesionidae, and Microdesmidae (lumped with Ophidiidae, Labridae, Lutjanidae, and Gobiidae, respectively). In addition, four families currently recognized as separate entities await formal nomenclatural description in compliance with the International Code of Zoological Nomenclature (ICZN) (i.e., "Cyclopsettidae", "Percalatidae", "Percophidae", "Rivulidae" and “Pantanodontidae”). The ordinal status of 32 percomorph families (vs. 50 in version 1belonging to the Series Carangimorpharia, Ovalentaria, and Eupercaria still remain uncertain (incertae sedis) due to either poor phylogenetic resolution or unavailability of genetic data. We therefore list these families as incertae sedis within each of these groups (Carangimorpharia, Ovalentaria, and Eupercaria) awaiting new phylogenetic evidence to clarify their ordinal status. Non-monophyletic families in this version (24 vs. 40 in version 1) are: Acropomatidae, Alepocephalidae, Bathydraconidae, Bathymasteridae, Chaenopsidae, Cheilodactylidae, Chlorophthalmidae, Clupeidae, Gempylidae, Grammatidae, Hemiramphidae, Ipnopidae, Labrisomidae, Macrouridae, Nototheniidae, Paralepididae, Phosichthyidae, Scombridae, Scopelarchidae, Scorpaenidae, Stichaeidae, Synodontidae, Trachichthyidae, Zenarchopteridae (see details below). 

This version includes comments about the recent classification of Nelson et al. (2016). For stability purposes, it incorporates several taxon names recognized by these authors, as long as they are based on monophyletic groups in our treeExamples include classification of suborders in Osmeriformes, Zeiformes and Beryciformes, validation of Trachichthyifomes and recognition of Acanthopterygii. A complete list of 26 changes made in accordance with Nelson et al. (2016) can be found here. We note, however, that many of the groups classified by Nelson et al. (2016) are incongruent with our phylogeny and thus are not recognized herein (though discrepancies are noted in each case). Examples of non-monophyletic taxa as circumscribed by Nelson et al. (2016) include Osmeromorpha, Scombriformes, Moroniformes and Perciformes. The tables at the bottom of the classification provide an exhaustive comparison of ordinal/supraordinal taxa (Table 1) or families (Table 2) that differ from Nelson et al. (2016). Table 2 also lists differences about the families recognized by van der Laan et al. (2014).

The new classification scheme presented here (version 4) 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 the authors; for the most updated version always visit DeepFin.

Version 4 presents new names. A complete list of changes from previous versions can be found here.

Figure 1. Fish Tree of Life. The backbone tree is from Betancur-R. et al. (2015), with two crown clades grafted into it: percomorphs (Sanciangco et al., 2016) and otophysans (Arcila et al., 2017). The complete tree includes 1770 species of bony fishes. Numbers in parenthesis indicate number of orders and families included in each major clade. 

Phylogenetic Classification of Bony Fishes (version 4)

Megaclass Osteichthyes (= extant Euteleostomi)

Superclass Actinopterygii (100%)

Class Cladistia (100%)

Order Polypteriformes


Class Actinopteri (100%)

Subclass Chondrostei (100%)

Order Acipenseriformes



Subclass Neopterygii (100%)

Infraclass Holostei (100%)

Order Amiiformes (= extant Halecomorphi)


Order Lepisosteiformes (= extant Ginglymodi) (100%)           


Infraclass Teleostei (100%)

Comment: Teleosteomorpha, Teleostei and Teleocephala (crown Teleostei) are all synonyms when only extant members are considered.

Megacohort Elopocephalai sensu Arratia (1999) (100%)

Cohort Elopomorpha (100%)

Order Elopiformes (100%)



Order Albuliformes (95%)


Order Notacanthiformes (92%)



Order Anguilliformes (100%)










Not examined: Chlopsidae, Cyematidae, Derichthyidae (including Colocongridae; e.g., Johnson et al. (2012)), 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 our tree; thus, no subordinal classification is proposed.

Megacohort Osteoglossocephalai (= Osteoglossocephala sensu Arratia (1999)) (100%)

Supercohort Osteoglossomorpha sensu Arratia (1999)

Comment: previous versions of the classification validated the supercohort Osteoglossocephala as well as the cohort Osteoglossomorpha, which were redundant in content. For simplicity and to avoid confusion  Osteoglossocephala sensu Arratia (1999) is the same as Osteoglossocephalai here and in previous versions, but not the same as Osteoglossocephala in previous versions  we now name this supercohort Osteoglossomorpha, but this change also means that the endings for the ranks cohort and supercohort are interchangeable.

Order Hiodontiformes (100%)


Order Osteoglossiformes (42%)




Osteoglossidae (includes Arapaima and Heterotis, formerly in Arapaimidae; e.g., Wilson and Murray (2008))


Supercohort Clupeocephala sensu Arratia (2010) (100%)

Cohort Otomorpha (= Otocephala, Ostarioclupeomorpha) (92%)

Subcohort Clupei (= Clupeomorpha) (100%)

Order Clupeiformes (100%)

Suborder Denticipitoidei


Suborder Clupeoidei (98%)


Clupeidae (not monophyletic)



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. The family Sundasalangidae no longer recognized because Sundasalanx is nested within Clupeidae (Lavoué et al., 2013)Clupeidae also includes the round herrings (subfamily Dussumieriinae; Lavoué et al. (2013)), sometimes placed in the family Dussumieriidae (van der Laan et al., 2014).

Subcohort Alepocephali (37%)

Order Alepocephaliformes

Alepocephalidae (not monophyletic)


Comment: Alepocephalidae includes Bathylaco, placed in Bathylaconidae by Nelson et al. (2016), and the former Leptochilichthyidae (see Lavoué et al. (2008) and Paulsen et al. (2009)). The position of alepocephaloids as the sister group to Ostariophyisi also was reported by Lavoué et al. (2008) and Paulsen et al. (2009).

Subcohort Ostariophysi (99%)

Section Anotophysa (= Anotophysi) (100%)

Order Gonorynchiformes



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

      Comment: Suborders in Gonorynchiformes are no longer recognized. See also Nelson et al. (2016).

Section Otophysa (= Otophysi) (100%)

Comment: Although most molecular studies (e.g., Nakatani et al., 2011; Chen et al., 2014a) are incongruent regarding otophysan interrelationships, our recent investigation of this question using genome-wide exon data in combination with topology tests (Arcila et al., 2017provides overwhelming support for the null morphological hypothesis of Fink and Fink (1981), which places characiforms sister to a clade including siluriforms plus gymnotiforms. Three otophysan superorders (Cypriniphysae, Characiphysae and Siluriphysae) are now recognized. Their taxonomic composition is similar to that originally proposed by Fink and Fink (1981), except that Characiphysae now contains a single order (Characiformes) following Nelson et al. (2016).

Superorder Cypriniphysae (92%)

Order Cypriniformes

Botiidae (following Chen et al. (2009))






Not examined: BalitoridaeGastromyzontidae (following Kottelat (2012)), Barbuccidae, Ellopostomatidae, Psilorhynchidae, Serpenticobitidae, Vaillantellidae.

Superorder Characiphysae (= Characiphysi) (100%)

Order Characiformes

Suborder Citharinoidei (100%)



Suborder Characoidei (100%)





















Not examined: Bryconidae.

Comment: Although characifom monophyly has been elusive from most molecular studies (e.g., Nakatani et al., 2011; Chen et al., 2014a), our recent phylogenomic study provides overwhelming support about the monophyly of the order (Arcila et al., 2017). 

Superorder Siluriphysae (= Siluriphysi) (100%)

Order Gymnotiformes (100%) 

Suborder Gymnotoidei


Suborder Sternopygoidei (not monophyletic here but see Tagliacollo et al. (2015))





Comment: Tagliacollo et al. (2015) proposed a revised classification for Gymnotiformes based on the most comprehensive phylogenetic analyses of the order to date, using both multi-locus sequence data and morphological evidence. They obtained two major clades within Sternopygoidei, which they named Rhamphichthyoidea (Rhamphichthyidae + Hypopomidae) and Sinusoidea (Sternopygidae + Apteronotidae). Although ranks for these clades are not explicit in their classification scheme, the endings suggest that these are superfamilies. According to the ICZN (article 61.2.2) “when a nominal taxon in the family group… is raised or lowered in rank, or its name is used at more than one rank simultaneously, the name-bearing type remains the same [Arts. 36.2, 43.1, 46.2].” In other words, the proper superfamily name for the “Sinusoidea” clade should be Sternopygoidea (suborder Sternopygoidei), to reflect a name-bearing type. Aside from the nomenclatural issues, our gymnotiform clade (based on genome-wide data; Arcila et al. (2017)) does not support the monophyly of "Sinusoidea."

Order Siluriformes (100%)

Suborder Loricarioidei (75%)






Not examined: Scoloplacidae.

Suborder Diplomystoidei


Suborder Siluroidei (100%)





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














Not_examined: AkysidaeAmphiliidaeAnchariidaeAriidaeAuchenoglanididaeAustroglanididae


Lacantuniidae, MalapteruridaeSchilbeidaeRitidae.

Comment: Recognition of catfish families follows Sullivan et al. (2006) and Lundberg et al. (2007), except for Ailidae, Auchenoglanididae and Ritidae that are herein recognized following Nelson et al. (2016), and Kryptoglanidae that follows Britz et al. (2014). The subordinal classification is based on Sullivan et al. (2006).

Supercohort Clupeocephala (cont.)

Cohort Euteleosteomorpha (= Euteleostei) (100%)

Subcohort Lepidogalaxii

Order Lepidogalaxiiformes


Subcohort Protacanthopterygii sedis mutabilis (100%)

Order Galaxiiformes (94%)


Order Argentiniformes (47%)





Order Salmoniformes (62%)


Order Esociformes (100%)



Comments: Nelson et al. (2016) restricted the circumscription of Protacanthopterygii to the clade including Salmoniformes and Esociformes, and also placed the orders Galaxiiformes and Argentiniformes, along with Stomiatiformes and Osmeriformes, in a new taxon they named Osmeromorpha. Circumscription of Osmeromorpha follow the results of the molecular phylogeny of Burridge et al. (2012). It is noteworthy that Burridge et al.'s study was designed to address intrafamilial galaxiid relationships. Their selection of non-galaxiid outgroups was only for time-calibration purposes; they did not intend to assess supraordinal relationships among early euteleosts. In addition to Osmeromorpha, Nelson et al. (2016) classified a purported clade including most euteleosts, except for Lepidogalaxiiformes, Salmoniformes and Esociformes, in an unranked taxon named Zoroteleostei by Wilson and Williams (2010). Circumscriptions of Osmeromorpha and Zoroteleostei  sensu Nelson et al. (2016) are incongruent with all recent higher-level phylogenetic analyses of fishes (e.g., Near et al., 2012a; Betancur-R. et al., 2013aBroughton, et al., 2013Betancur-R. et al., 2015).

Subcohort Stomiati (100%) (see comments under Protacanthopterygii above)

Order Stomiatiformes sensu Rosen (1973) (= Stomiiformes sensu Fink and Weitzmann (1982)) (100%)

Suborder Gonostomatoidei (54%)

Comment: Diplophidae is no longer recognized as a separate family; it is listed as subfamily of Gonostomatidae in van der Laan et al. (2014) and Nelson et al. (2016)Diplophos is sister to all other gonostomatids in the tree. 

Suborder Stomiatoidei (=Phosichthyoidei) (61%)

Comments: Suborders in Stomiatiformes are now recognized following Nelson et al. (2016), except that their Phosichthyoidei is named Stomiatoidei herein (based on Stomiidae).

Order Osmeriformes (100%)

Suborder Osmeroidei (100%)

Suborder Retropinnoidei (100%)

Retropinnidae (including Prototroctidae following Nelson et al. (2016))

Comment: suborders in Osmeriformes are now classified following Nelson et al. (2016).

Subcohort Neoteleostei (100%)

Infracohort Ateleopodia (= Ateleopodomorpha) (98%)

Order Ateleopodiformes


Infracohort Eurypterygia (=Eurypterygii) (96%)

Section Cyclosquamata (= Aulopa) (100%)

Comment: We now recognize Cyclosquamata sensu Rosen following other recent classifications (e.g., Davis (2010)Nelson et al. (2016); = Aulopa in Wiley and Johnson (2010) and in previous versions of this classification).

Order Aulopiformes

Suborder Aulopoidei (not monophyletic)



Synodontidae (not monophyletic)

Suborder Paraulopoidei


Suborder Alepisauroidei (not monophyletic)

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


Chlorophthalmidae (not monophyletic)



Ipnopidae (not monophyletic)



Paralepididae (not monophyletic)

Scopelarchidae (not monophyletic)

Sudidae (following Davis (2010); synonym of Paralepididae according to van der Laan et al. (2014))

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

Comments: Anotopteridae is not recognized (lumped with Alepisauridae); aulopiform families listed follow Davis (2010). Lestidiidae is now recognized following Ghedotti et al. (2014) and Nelson et al. (2016). Although not monophyletic herein, the monophyly of aulopiform suborders is supported by Davis (2010).

Section Ctenosquamata (97%)

Subsection Myctophata (= Scopelomorpha) (100%)

Order Myctophiformes



Subsection Acanthomorphata (= Acanthomorpha) (96%)

Division Lampripterygii (= Lampridacea in previous versions; = Lamprimorpha in Nelson et al. (2016)(82%)

Comment: Endings for the rank "division" have been changed to "-pterygii" (see comments under Acanthopterygii below). 

Order Lampriformes (= Lampridiformes in previous versions, = Allotriognathi)

Lampridae (= Lamprididae in previous versions)




Not examined: Radiicephalidae, Veliferidae.

Division Paracanthopterygii (sensu Miya et al. (2005) and Grande et al. (2013) but excluding Polymixiidae; = Paracanthomorphacea in previous versions(100%)

Comment: Endings for the rank Division have been changed to "-pterygii" (see comments under Acanthopterygii below). 

Series Percopsaria (100%)

Order Percopsiformes




Series Zeiogadaria (= Zeiogadiformes sensu Li et al. (2009)) (100%)

Subseries Zeiariae (100%)

Order Zeiformes

Suborder Cyttoidei

 Not examined: Cyttidae.

Suborder Zeiodei

Not examined: GrammicolepididaeOreosomatidae.

Comment: Zeiform suborders are now classified following Tyler (2003) and Nelson et al. (2016).

Subseries Gadariae (100%)

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


Order Gadiformes (100%)

Suborder Macrouroidei (100%)

Macrouridae (not monophyletic here, but see Roa-Varón and Ortí (2009); includes taxa often placed in Bathygadidae and Trachyrincidae)

Steindachneriidae (following Roa-Varón and Ortí (2009); synonym of Merluciidae according to van der Laan et al. (2014))

Suborder Gadoidei (not monophyletic)

Gadidae (includes taxa often placed in Ranicipitidae)

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


Merlucciidae (includes taxa often placed in Macruronidae)



Not examined: Bregmacerotidae, Euclichthyidae, Melanonidae.

Suborder Muraenolepidoidei


Comment: The classification of suborders and families in Gadiformes follows Roa-Varón and Ortí (2009: fig. 6). There are many other classification proposals for the order, however, that are incongruent (see Roa-Varón and Ortí, 2009: table 1).

Division Polymixiipterygii (100%)

Comment: Endings for the rank Division have been changed to "-pterygii" (see comments under Acanthopterygii below). 

Order Polymixiiformes


Comment: We place Polymixiidae in its own division (as opposed to Paracanthopterygii as in Miya et al. (2005) and Grande et al. (2013)) to recognize its rogue placement among early acanthomorph lineagesThis classification system is not only congruent with our tree but also robust to phylogenetic uncertainty.

Division Acanthopterygii (= Euacanthomorphacea in previous versions) (95%)

Comment: Previous versions of this classification named this clade Euacanthomorphacea, a taxon recognized by Johnson and Patterson (1993) to included polymixiids, percopsids and crown acanthomorphs. Because polymixiids and percopsids are not members of this group, it seems reasonable to instead adopt Acanthopterygii, recognizing its extensive use in ichthyology. Note that Acanthopterygii was not classified by Wiley and Johnson (2010). This change follows Near et al. (2013) and Nelson et al. (2016). For consistency, we also changed all Division suffixes to "-pterygii."

Subdivision Berycimorphaceae (100%)

Order Beryciformes (100%)

Suborder Berycoidei (100%)

Suborder Stephanoberycoidei (78%)

Comment: Beryciform suborders are now classified following Nelson et al. (2016).

Order Trachichthyiformes sensu Moore (1993) (100%)





Trachichthyidae (not monophyletic)

Comments: Beryciformes sensu lato (as in previous versions) is now split into Beryciformes sensu stricto (including suborders Berycoidei and Stephanoberycoideiand Trachichthyiformes sensu Moore (1993), following Nelson et al. (2016). The subordinal classification for Trachichthyiformes proposed by Nelson et al. (2016), however, is incongruent with the tree and therefore not is implemented herein. 

Subdivision Holocentrimorphaceae (100%)

Order Holocentriformes


Comment: Stiassny and Moore (1992) and Moore (1993) provide morphological evidence supporting a sister-group relationship between holocentrids and percomorphs, validating the 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 (100%)

Ophidiidae (includes the former Carapidae)

Suborder Bythitoidei (100%)


Bythitidae (not monophyletic)

Not examined: Parabrotulidae.

Comments: Due to phylogenetic nestedness, we now synonymize Carapidae with Ophidiidae. Recognition of Dinematichthyidae follows Møller et al. (2016); raised from subfamily Dinematichthyinae (formerly Bythitidae). These authors also lumped Aphyonidae with Bythitidae; thus Aphyonidae is no longer validated.

Series Batrachoidaria (100%)

Order Batrachoidiformes


Series Syngnatharia (84%)         

Order Syngnathiformes

Suborder-level incertae sedis in Syngnathiformes


Centriscidae (including taxa often placed in Macroramphosidae)



Suborder Syngnathoidei (100%)


Not examined: Solenostomidae (assumed affinity with Syngnathidae).

Suborder Dactylopteroidei (100%)


Suborder Callionymoidei (= Callionymiformes sensu Nelson et al. (2016)) (100%)


Not examined: Draconettidae (assumed affinity with Callionymidae).

Suborder Mulloidei (100%)


Series Pelagiaria (= Stromateoidei sensu Li et al. (2009); = Pelagia sensu Miya et al. (2013), = Scombrimorpharia in previous versions) (99%)          

Order Scombriformes







Gempylidae (not monophyletic)




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




Not examined: Amarsipidae, Scombropidae, Tetragonuridae (Doiuchi et al., 2004; Miya et al., 2013).

Comment: Interfamilial resolution in Scombriformes is tenuous; classification of scombriform families into suborders (e.g., Scombroidei, Stromateoidei, Icostoidei) or new orders requires further work. Our circumscription of Scombriformes includes taxa placed by Nelson et al. (2016) in the orders Scombriformes, Trachiniformes in part, Icosteiformes and Scombrolabraciformes.

Series Gobiaria (= Gobiiformes sensu Thacker (2009) and Thacker et al. (2015)(100%)

Order Kurtiformes (= Apogonoidei sensu Thacker et al. (2015)(100%)

Suborder Kurtoidei (100%)


Suborder Apogonoidei (100%)


Comment: Johnson (1993) noted that the configuration of dorsal gill-arch elements may be homologous in Kurtus and apogonids.

Order Gobiiformes (100%)

Suborder Trichonotoidei

Not examined: Trichonotidae.

Suborder Gobioidei (100%)


Eleotridae (includes the former Xenisthmidae; see Thacker (2009))

Gobiidae (includes the former MicrodesmidaeKraemeriidae, Ptereleotridae, and Schindleriidae; see Thacker (2009) and Thacker et al. (2015))


Not examined: Oxudercidae (= Gobionellidae), Milyeringidae, Rhyacichthyidae, Thalasseleotrididae.

Comment: Recognition of Butidae, Oxudercidae and Milyeringidae follows Thacker (2009) and Thacker et al. (2015). We now recognize Oxudercidae instead of Gobionellidae (Gobionellidae is a junior synonym).

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)The classification of suborders in Gobiiformes now follows Thacker et al. (2015), but with modifications. Our circumscription of Kurtiformes is the same as Apogonoidei in Thacker et al. (2015). These authors also identified Trichonotus as the sister lineage of the gobies; thus, we place Trichonotus in its own suborder (Trichonotoidei). Finally, Odontobutoidei and Eleotroidei, validated in previous versions of the classification, are now considered synonyms of Gobioidei. 

Series Anabantaria (= Anabantiformes sensu Li et al. (2009)) (100%)

Order Synbranchiformes (100%)

Suborder Mastacembeloidei (100%)


Not examined: Chaudhuriidae.

Suborder Indostomoidei (100%)


Suborder Synbranchoidei


Order Anabantiformes (= Labyrinthici) (100%)

Suborder Anabantoidei (100%)




Suborder Channoidei (100%)


Suborder Nandoidei (91%)




Comment: Suborders of Anabantiformes reflect well-supported and robust clades, but only when the family Channidae is placed in a separate suborder (Channoidei). Affinities of Channidae with other anabantiform families vary among studies (e.g., Near et al., 2013; Betancur-R. et al., 2013a; Sanciangco et al., 2016). The scheme presented with three suborders is robust to this ambiguity.

Series Carangaria (= Carangimorpha sensu Li et al. (2009)) (99%)

Order-level incertae sedis in Carangaria

Centropomidae (includes the former Latidae, following Greenwood (1976) and Li et al. (2011))







Order Istiophoriformes (100%)



Comment: Our tree does not support placement of Sphyraenidae in this order, as suggested by Nelson et al. (2016).

Order Carangiformes sedis mutabilis (not monophyletic)






Comment: Monophyly of Carangiformes is not significantly rejected by the data (Betancur-R. et al., 2013b).

Order Pleuronectiformes (21%)

Suborder Psettodoidei (100%)


Suborder Pleuronectoidei (99%)





"Cyclopsettidae" (awaits formal description)



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

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




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

Comment: Although contentious, the monophyly of Pleuronectiformes is resolved by several molecular studies (Betancur-R. et al., 2013b; Betancur-R and Ortí, 2014; Sanciangco et al., 2016). Paralichthyidae is monophyletic if the Cyclopsetta group is included in its own family (Betancur-R. et al., 2013b). Formal description of a new family for Cyclopsetta is needed in compliance with the ICZN (hence "Cyclopsettidae")

Series Ovalentaria (sensu Smith and Near in Wainwright et al. (2012); = Stiassnyiformes sensu Li et al. (2009)) (97%)

Order-level incertae sedis in Ovalentaria

Ambassidae (= Chandidae)

Congrogadidae sensu Godkin and Winterbottom (1985); formerly a subfamily of Pseudochromidae


Grammatidae (= Grammidae; not monophyletic here, but see Gill and Mooi (1993))






Superorder Cichlomorphae (40%)

Order Cichliformes



Comment: The circumscription of Cichliformes is expanded herein to include Pholidichthyidae (formerly Pholidichthyiformes; see Nelson et al., 2016).

Superoder Atherinomorphae (= Atherinomorpha) (100%)

Order Atheriniformes (100%)

Suborder Atherinopsoidei (100%)

Comment: Atherinopsidae includes the subfamilies Atherinopsinae, Notocheirinae and Menidiinae. The circumscription of Atherinopsidae sensu Nelson et al. (2016) includes only Menidiinae and Atherinopsinae, which renders Atherinopsidae non-monophyletic (Notocheirinae is nested within; see Campanella et al. (2015)).

Suborder Atherinoidei (100%)


Not examined: Dentatherinidae.

Comment: Classification of suborders and families in Atheriniformes follows Campanella et al. (2015)Notocheiridae is no longer recognized (subfamily of Atherinopsidae).

Order Beloniformes (79%)

Suborder Adrianichthyoidei


Suborder Belonoidei (100%) (= Exocoetoidei sensu Nelson et al. (2016))

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


Hemiramphidae (not monophyletic)

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

Order Cyprinodontiformes (100%)

Suborder Aplocheiloidei


Not examined: Nothobranchiidae, "Rivulidae."

Comment: According to van der Laan et al. (2014) the name Rivulidae Myers 1925 is preoccupied by Rivulini Grote 1895 in Lepidoptera (hence "Rivulidae").

Suborder Cyprinodontoidei (100%)




Not examined: Anablepidae, Goodeidae, Profundulidae, Valenciidae.

Possibly included: “Pantanodontidae” (requires formal description).

Comments: Cyprinodontidae and Poeciliidae are monophyletic here with reduced taxonomic sampling but not in two other recent studies that included a much broader sampling (Pollux et al., 2014Pohl et al., 2015). Pohl et al. (2015) identified a rogue placement for Pantanodon among cyprinodontiforms. The topology most often obtained by these authors included Pantanodon as sister to all cyprinodontoids. Formal description of a new family for Pantanodon is needed in compliance with the ICZN. 

Superorder Mugilomorphae (100%)

Order Mugiliformes


Superorder Blenniimorphae (90%)

Order Gobiesociformes (100%)


Comment: The order Gobiesociformes is recognized herein following Wiley and Johnson (2010) and Nelson et al. (2016).

Order Blenniiformes (39%)


Chaenopsidae (not monophyletic)



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


            Comments: Circumscription of Blenniiformes follows Lin and Hastings (2013). The reciprocal monophyly of gobiesocoids and blennioids is supported by molecular (Wainwright et al., 2012Lin and Hastings; 2013) and morphological (Springer and Orrell, 2004) evidence. Our new tree also resolves the blennioids as monophyletic, a result not obtained in our previous large-scale analyses. According to Lin and Hastings (2013)Chaenopsidae is monophyletic if Stathmonotus is included in Labrisomidae

Series Eupercaria (83%)

Order-level incertae sedis in Eupercaria












Not examined: Dinolestidae, Dinopercidae (see Smith and Craig; 2007), Parascorpididae 

Comment: The family Parascorpididae, traditionally classified in “Perciformes”, is provisionally listed here; it is not placed in the recently circumscribed Perciformes given the long history of phylogenetic indistinctiveness between Percoidei, Perciformes, and Percomorpha (Smith and Craig, 2007). 

Order Gerreiformes (100%)


Comment: Validation of Gerreiformes (resurrected herein) reflects the placement of Gerreidae as sister to all other eupercarians.

Order Uranoscopiformes (= Paratrachinoidei sensu Li et al. (2009)) (98%)


Cheimarrichthyidae (= Cheimarrhichthyidae)



Order Labriformes (100%)

Labridae (includes taxa previoulsy listed in Scaridae and Odacidae; see also Wainwright et al. (2012))

Order Ephippiformes (100%)



Comments: Greenwood et al. (1966) hypothesized a close affinity between Drepane and ephippids. Nelson et al. (2016) named this clade Moroniformes, including Moronidae in addition to Drepaneidae and Ephippidae. Our results do not support the placement of Moronidae in this order.

Order Chaetodontiformes (66%)



Comment: This clade has been consistently obtained by previous studies with higher nodal support.

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




Comments: Nelson et al. (2016) included Emmelichthyidae and Sciaenidae in this order, in addition to AcanthuridaeLuvaridae and Zanclidae. Our results do not support the placement of Emmelichthyidae and Sciaenidae in Acanthuriformes.

Order Lutjaniformes, new circumscription (59%)


Lutjanidae (includes the former Caesionidaer; e.g., Johnson (1993) and Miller and Cribb (2007))

Comments: The order Lutjaniformes (a Bleeker name) is herein resurrected for the clade including lutjanids and haemulids. Although nodal support is low, this clade is often obtained in various large-scale studies.

Order Lobotiformes (100%)

Hapalogenyidae (= Hapalogeniidae)



Comment: See Sanciangco et al. (2016) for a discussion on the circumscription and morphological support of Lobotiformes. 

Order Spariformes sensu Akazaki (1962) and Johnson (1981) (87%)



Sparidae (includes the former Centracanthidae)

Comments: 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. The family Centracanthidae is no longer recognized as valid; synonym of Sparidae following Santini et al. (2014) and Sanciangco et al. (2016)Nelson et al. (2016) also included in this order the families CallanthiidaeLobotidae (including Datnioididae) and Sillaginidae. Our results do not support the placement of these three or four families in Spariformes.

Order Priacanthiformes, new circumscription (98%)



Comments: A sister-group relationship between cepolids and priacanthids is strongly supported by other molecular studies as well as by larval morphology (Leis and Carson-Ewart, 2000). Priacanthiformes is a Bleeker name.

Order Caproiformes (37%)


Comment: This order is herein recognized following Nelson et al. (2016).

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%)


Suborder Antennarioidei (100%)


Not examined: Brachionichthyidae, Lophichthyidae, Tetrabrachiidae.

Suborder Chaunacoidei (100%)


Suborder Ogcocephaloidei (100%)


Suborder Ceratioidei (100%)






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

Order Tetraodontiformes (100%). This order is the sister group of Lophiiformes (see comments under Lophiiformes above).

Suborder Triodontoidei


Suborder Triacanthoidei


Suborder Triacanthodoidei (100%) 


Suborder Tetraodontoidei (100%)



Suborder Moloidei (100%)


Suborder Balistoidei (100%)



Suborder Ostracioidei (100%)



Comments: This subordinal classification differs from that proposed by Santini and Tyler (2003). It is robust to phylogenetic uncertainty and has recently been adopted by other (e.g., Bannikov et al., 2016)

Order Pempheriformes (33%)

Acropomatidae (not monophyletic)













"Percophidae" (see comments)



Not examined: Leptoscopidae (assumed affinity with Creediidae; see Odani and Imamura (2011)).

Comments: 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. Because Percophis brasiliensis (type species of Percophidae) is a Notothenioid (Near el al., 2015), and the remaining “percophids” are in Pempheriformes, then the pempheriform “percophids” require family relocation. The subfamily Hemerocoetinae Kaup 1873 is raised herein to the family level, following Thacker et al. (2015): “Additional proposed changes to the classification of Percomorpha include... recognition of Hemerocoetidae as a taxonomic family containing Matsubaraea, Enigmapercis, PteropsaronAcanthaphritis, and Osopsaron and the unsampled Dactylopsaron, Hemerocoetes, and Squamicreedia.

Order Centrarchiformes (98%)

Suborder Centrarchoidei (93%)




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

Comment: Inclusion of Enoploside in this suborder differs from the results obtained by Lavoué et al. (2014).

Suborder Cirrhitoidei (similar to Cirrhitoidea sensu Greenwood (1995) and Burridge and Smolenski (2004); treated as Cirrhitiformes in previous versions of the classification) (97%)


Cheilodactylidae (not monophyletic)




Comment: The families LatridaeChironemidae and Aplodactylidae are nested within Cheilodactylidae, rendering the latter non-monophyletic (Sanciangco et al., 2016).

Suborder Percichthyoidei (100%)

Percichthyidae (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 (2016), but the type species of Macquaria (M. australasica) is closely related to other species of Macquaria (M. ambigua) within Percichthyidae sensu stricto, thus both are valid genus names (Peter Unmack et al., pers. comm.; Lavoué et al. (2014)). Percichthyidae sensu stricto includes Percilia (formerly placed in its own family Perciliidae).

Suborder Percalatoidei (100%)

"Percalatidae" (awaits formal description)

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); = Terapontiformes in previous versions of the classification) (99%)


Girellidae (following Carpenter (2001); listed as a subfamily of Kiphosidae by van der Laan et al. (2014))





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), Chen et al. (2014b) and Lavoué et al. (2014).

Order Perciformes (= Serraniformes sensu Li et al. (2009) and Lautredou et al. (2013); including taxa previously listed in Scorpaeniformes, Cottiformes, Gasterosteiformes and Trachiniformes) (93%)

Suborder Bembropoidei, new (100%)


Comment: This suborder is newly classified to accommodate the family Bembropidae. Bembropidae is recognized following Smith and Craig (2007); it is a synonym of Percophidae according to van der Laan et al. (2014).

Suborder Normanichthyoidei

            Not examined: Normanichthyidae.

Comment: This suborder is classified following van der Laan et al. (2014) and Nelson et al. (2016).

Suborder Serranoidei sedis mutabilis (64%)


Comment: We do not recognize Epinephelidae as a separate family, following Smith and Craig (2007) and Ma et al. (2016). The main justification for such nomenclatural change was that Smith and Craig's phylogenetic analysis failed to resolve the monophyly of serranids (including epinephelines, anthiines and serranines); however, they did not conduct a topology test to ask whether this null hypothesis is rejected by their data. Our large-scale tree supports the monophyly of Serranidae, albeit with low support. Also, while elevating a subfamily to a family is a minor nomenclatural change, this rearrangement creates confusion for fish managers and conservation biologists given the commercial importance of groupers and the endangered status of many species.

Suborder Percoidei, restricted circumscription (99%)

            Niphonidae (following Smith and Craig (2007); synonym of Serranidae according to van der Laan et al. (2014))


            Not examined: Trachinidae.

Comment: Lautredou et al. (2013) obtained a clade uniting Percidae and Trachinidae with full support, based on the analysis of seven nuclear markers .

Suborder Notothenioidei (100%)


Bathydraconidae (not monophyletic)





Nototheniidae (not monophyletic)



Comment: Percophidae is herein placed in Notothenioidei following Near el al. (2015); see comments above under Pempheriformes.

Suborder Scorpaenoidei (72%)

            Scorpaenidae (not monophyletic)





Not examined: ApistidaeAploactinidaeCongiopodidaeEschmeyeridaeGnathanacanthidaeNeosebastidae

PataecidaePerryenidae (sensu Honma et al., 2013) and Zanclorhynchidae.

Comment: Nine families now included in Scorpaenoidei were listed in previous versions of this classification as not examined under Perciformes. See also Imamura (2004).

Suborder Platycephaloidei (= Bembroidei in previous versions) (26%)





Not examined: Plectrogeniidae (see Imamura, 2004).

Comment: Previous versions of this classification included Bembridae and Parabembridae in the suborder Bembroidei, which we now expand to also include HoplichthyidaePlatycephalidae and Plectrogeniidae (previously listed as suborder-level incertae sedis in Perciformes) – a well supported clade in our tree (100% BS). We now name this taxon Platycephaloidei in accordance to other classifications (e.g., Imamura, 1996). Note that the family composition differs from that in other studies as Peristediidae and Triglidae are placed in a different suborder (Triglioidei).

Suborder Triglioidei sensu Jordan (1923) (100%)



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 possibly sister groups (although not in our results); they have been grouped in a clade named Zoarciformes by Li et al. (2009).

Infraorder Anoplopomatales (= Anoplopomatoidei in previous classifications)


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


Bathymasteridae (not monophyletic)



Stichaeidae (not monophyletic)



Not examined: Eulophiidae (sensu Kwun and Kim (2013); see also Nelson et al. (2016)), PtilichthyidaeScytalinidae.

Infraorder Gasterosteales (similar Gasterosteoidei in other classifications, but excluding Indostomidae) (100%)




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

Zaniolepididae (sensu Smith and Busby (2014), formerly a subfamily of Hexagrammidae (van der Laan et al., 2014))

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 Zaniolepididae 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 Cottales (99%) (= Cottoidei sensu Smith and Busby (2014))   

Agonidae (includes the former Hemitripteridae; see Smith and Busby (2014))

Cottidae (includes the former Abyssocottidae, Comephoridae, and Cottocomephoridae; see Smith and Busby (2014))



Psychrolutidae (includes the former Bathylutichthyidae and many marine genera previously placed in Cottidae; see Smith and Busby (2014))

Scorpaenichthyidae (following  Smith and Busby (2014))


Not examined: Jordaniidae (sensu Smith and Busby (2014)) and Rhamphocottidae (including the former EreuniidaeSmith and Busby (2014)).

Comment: Smith and Busby (2014) changed the membership of Agonidae, Cottidae and Psycholutridae to achieve monopyhyly of these families; our phylogenetic results support their revised circumscription.

Superclass Sarcopterygii (58%)

Class Coelacanthimorpha (= Actinistia)

                  Order Coelacanthiformes


Class Dipnotetrapodomorpha (100%)

Subclass Dipnomorpha

        Superorder Ceratodontae (= Dipnoi)

      Order Ceratodontiformes

                  Suborder Ceratodontoidei


                  Suborder Lepidosirenoidei (100%)



                  Subclass Tetrapodomorpha (100%)

Comment: Recent genomic evidence supports the sister-group relationship between lungfishes and tetrapods (Chen et al., 2015; Irisarri and Meyer, 2016).

Table 1. Remarkable differences for ordinal or supraordinal taxa between Nelson's et al. (2016) classification and the update proposed herein. The circumscription of other orders may also differ due to variations in family   validation (see Table 2) or due to inclusion of fossil taxa in Nelson's et al. (2016). Differences in taxonomic ranks and endings are considered minor and not listed herein.

Taxon (order-level or above)

Differences with NGW


Teleostomi, Ginglymodi, Halecomorphi and Teleosteomorpha/Teleocephala

Not classified herein

Redundant with Osteichthyes, Amiiformes, Lepisosteiformes, and Teleostei, respectively, when only extant taxa are considered.


Not classified by NGW

Only shown in one of NGW's cladograms but not formally classified therein.


Not classified by NGW

Non-polypteriform actinopterygiians.


Not classified by NGW

Not a major difference; it is redundant with Elopomorpha.


Classified into suborders in NGW but not herein

Phylogenetic incongruence with traditional subordinal classifications.


Not classified herein

The subordinal classification for Siluriformes follows Sullivan et al. (2006).


Includes four orders herein and only two in NGW

Differences are in part due to phylogenetic uncertainty. We classify this taxon as sedis mutabilis.


Classified by NGW only

Circumscription of this taxon is in conflict with Protacanthopterygii. See comments in text.


Classified by NGW only

Circumscription of this taxon is incongruent with all recent higher-level phylogenetic analyses of fishes. See comments in text.


Not classified by NGW

The circumscription of Stomiati herein is in conflict with NGW's Osmeromorpha. See comments above and in text.



Stomiatiformes sensu Rosen (1973); Stomiiformes sensu Fink and Weitzmann (1982)


Phosichthyoidei sensu NGW; Stomiatoidei herein

Based on Stomiidae


Includes Polymixiiformes in NGW but not here

Polymixia has a rogue placement among early acanthomorphs. Our classification is robust to phylogenetic uncertainty.


Not classified by NGW

Denotes a robust clade including Zeiformes + (Stylephoriformes + Gadiformes); this taxon has been recognized before (i.e., Zeiogadiformes sensu Li et al. (2009)).


Not classified herein

Subordinal classification of Gadiformes follows Roa-Varón and Ortí (2009) herein


Not classified by NGW

Subordinal classification of Gadiformes follows Roa-Varón and Ortí (2009) herein


Berycida sensu NGW is similar to Berycimorphaceae as classified herein, but the former includes Holocentriformes

Holocentridae is sometimes recovered as the sister taxon of percomophs, which may render Berycida sensu NGW not monophyletic.


Not classified herein

Not monophyletic


Not classified herein

Not monophyletic


Not classified by NGW

Included in Berycida sensu NGW. See comments above.


Not classified by NGW

A robust calde (series) including 17 families in the order Scombriformes, as classified herein.


Includes 17 families herein and only nine in NGW

Scombriformes sensu NGW is paraphyletic in light of all higher-level molecular phylogenies of percomorphs.

Scombroidei and Stromateoidei

Not classified herein

Interfamilial resolution in Scombriformes is tenuous; classification of scombriform families into suborders requires further work.


Not classified herein

Icosteidae, the sole family in this order, is part of Pelagiaria (Scombriformes) herein.


Not classified herein

Scombrolabracidae, the sole family in this order, is part of Pelagiaria (Scombriformes) herein.


Not classified herein (similar to Uranoscopiformes)

Trachiniformes sensu NGW is polyphyletic. It includes families placed in Pelagiaria and Eupercaria.


Not classified by NGW

A robust clade (series) including 10 families in the order Syngnathiformes, as classified herein.


Includes 10 families herein and eight in NGW

Exclusion of Mullidae and Callionymoidei renders Syngnathiformes paraphyletic.


Not classified herein

Not monophyletic.


Suborder (Callionymoidei) of Syngnathiformes herein

Recognition of Callionymiformes as a separate order renders Syngnathiformes paraphyletic.


Not classified by NGW

A robust clade (series) including the orders Synbranchiformes (including Indostomidae) and Anabantiformes.


Not classified by NGW

Indostomidae is not included in Synbranchiformes by NGW; exclusion of this family renders the order (and its component suborders) not monophyletic.


Not classified by NGW

The order Anabantiformes in classified in three suborders herein. This scheme is robust to phylogenetic uncertainity.


Not classified by NGW

A robust clade (series) including the orders Istiophoriformes, Carangiformes, Pleuronectiformes and several families listed as order-level incertae sedis.


Includes two families herein and three in NGW

Inclusion of Sphyraenidae renders Istiophoriformes non-monophyletic.


Exocoetoidei sensu NGW; Belonoidei herein

Belonoidei is the name-bearer


Not classified by NGW

A robust clade (series) including at least 16 orders and 161 families. The order Perciformes is in this clade.


Not classified by NGW

Includes the family Gerreidae, which is the sister to all other eupercarians (listed under Perciformes in NGW).


Includes three families herein and five in NGW

Inclusion of Emmelichthyidae and Sciaenidae renders Acanthuriformes non-monophyletic.

Acanthuroidei and Sciaenoidei

Not classified herein

See comment under Acanthuriformes above


Moroniformes in NGW (three families) and Ephippiformes herein (two families)

Our results do not support a close relationship between Moronidae and Drepaneidae + Ephippidae.


Includes three families herein and six in NGW

Inclusion of Callanthiidae, Lobotidae (including Datnioididae) and Sillaginidae renders Spariformes non-monophyletic.


Not classified by NGW

A robust clade (order) including the families Chaetodontidae and Leiognathidae.


Not classified by NGW

A robust clade (order) including the families Hapalogenyidae, Datnioididae and Lobotidae (listed in Spariformes or Perciformes in NGW).


Not classified by NGW

A robust clade (order) including the families Lutjanidae and Haemulidae (listed under Perciformes in NGW).


Not classified by NGW

A robust clade (order) including the families Priacanthidae and Cepolidae (listed under Perciformes in NGW).


Not classified by NGW

A robust clade (order) including the families Ammodytidae, Cheimarrichthyidae, Pinguipedidae and Uranoscopidae (listed under Trachiniformes in NGW).


Not classified by NGW

Placement of Molidae in Tetraodontoidei often results in the non-monophyly of Tetraodontoidei. The subordinal classification for Tetraodintiformes is robust to phylogenetic uncertainty


Not classified by NGW

Placement of Triacanthodidae in Triacanthoidei often results in the non-monophyly of Triacanthoidei. The subordinal classification for Tetraodintiformes is robust to phylogenetic uncertainty



NGW recognize the "Suborder Ostracioidea", but the correct ending for the rank suborder is "-iodei."


Not classified by NGW

A robust clade (order) including 17 families (most listed under Perciformes in NGW).


Not classified by NGW

A robust clade (order) including five suborders and 18 families (most listed under Perciformes in NGW).


Includes 61 families herein and 62 in NGW (but with very different circumscriptions)

Our definition of Perciformes is monophyletic; NGW maintain the status quo by using Perciformes as a taxonomic waste basket (polyphyletic).


Includes three families herein and 46 in NGW

Our definition of Percoidei is monophyletic; NGW maintain the status quo by using Percoidei as a taxonomic waste basket (polyphyletic).


Not classified by NGW

Includes Serranidae only.


Not classified by NGW

Includes Bembropidae.


Includes nine families herein and eight in NGW

The family Percophidae is a member of Notothenioidei herein (following Thacker et al. (2015)), weheras in NGW it belongs in Trachiniformes.


Includes several families that are part of four different perciform suborders herein

Recognition of Scorpaeniformes as a separate order renders Perciformes non-monophyletic.


Gasterosteoidei in NGW and Gasterosteales herein

Recognition of Scorpaeniformes as a separate order renders Perciformes non-monophyletic.


Gasterosteoidei in NGW and Gasterosteales herein

Gasterosteales herein is similar to Gasterosteoidei sensu NGW, except that the former excludes Indostomidae (classified under Synbranchiformes herein)

NGW: Nelson et al. (2016)

Table 2. Differences in the recognition of families between Nelson's et al. (2016) and van der Laan et al. (2014) classifications and the update proposed herein. Taxa are listed in alphabetic order.


Differences with NGW

Differences with vdLE



Provisionally recognized as "Cyclopsettidae" herein

Provisionally recognized as "Cyclopsettidae" herein

Awaits formal description; see Betancur-R. et al. (2013b)


Provisionally recognized as "Pantanodontidae" herein

Provisionally recognized as "Pantanodontidae" herein

Awaits formal description; see Pohl et al. (2015)


Provisionally recognized as "Percalatidae" herein

Provisionally recognized as "Percalatidae" herein

Awaits formal description; see text


Provisionally recognized as "Percophidae" herein

Provisionally recognized as "Percophidae" herein

Awaits formal description; lineage in Pempheriformes not related to Percophidae (Perciformes); see also Thacker et al. (2015)


Provisionally recognized as "Rivulidae" herein; Rivulidae in NGW


The name Rivulidae is preoccupied in Lepidoptera (see vdLE)



Synonym of Cottidae herein

Following Smith and Busby (2014)


Synonym of Rhombosoleidae herein

Synonym of Rhombosoleidae herein

Lumped due to phylogenetic nestedness (e.g., Betancur-R. et al., 2013b)


Synonym of Paralepididae in NGW


Following Davis (2010)


Synonym of Bythitidae herein


Following Møller et al. (2016)


Subfamily of Scorpaenidae in NGW


Following vdLE and Imamura (2004)



Synomym of Osteoglossidae herein

Following Wilson and Murray (2008)


Synonym (subfamily) of Atherinopsidae herein


Following Campanella et al. (2015)


Synonym (subfamily) of Macrouridae herein


Following Roa-Varón and Ortí (2009)


Synonym of Alepocephalidae herein


Following vdLE


Synonym of Psychrolutidae herein

Synonym of Psychrolutidae herein

Following Smith and Busby (2014)


Subfamily of Ipnopidae in NGW


Following Davis (2010)


Subfamily of Melanotaeniidae in NGW


Following Campanella et al. (2015) and vdLE


Subfamily of Percophidae in NGW

Subfamily of Percophidae in vdLE

Following Smith and Craig (2007)



Subfamily of Cobitidae in vdLE

Following Chen et al. (2009)


Subfamily of Characidae in NGW


Following Arcila et al. (2017) and vdLE



Subfamily of Eleotridae in vdLE

Following Thacker (2009) and Thacker et al. (2015)


Synonym of Lutjanidae herein

Synonym of Lutjanidae herein

Lumped due to phylogenetic nestedness


Synonym of Ophidiidae herein

Synonym of Ophidiidae herein

Lumped due to phylogenetic nestedness



Synonym of Sparidae herein

Following Santini et al. (2014) and Sanciangco et al. (2016)


Omitted by NGW and NEL


Following Arcila et al. (2017) and vdLE


Spelled Cheimarrhichthyidae in NGW


Following vdLE


Synomym of Derichthyidae in NGW

Synomym of Derichthyidae herein

Following Johnson et al. (2012)



Synonym of Cottidae herein

Following Smith and Busby (2014)


Subfamily of Pseudochromidae in NGW

Subfamily of Pseudochromidae in vdLE

Following our results and Godkin and Winterbottom (1985)



Synonym of Cottidae herein

Following Smith and Busby (2014)


Synonym of Lobotidae in NGW


Following Sanciangco et al. (2016)


Synonym of Brosmophycinae, a subfamily of Bythitidae in NGW


Following Møller et al. (2016)



Synonym of Clupeidae herein

Following Lavoué et al. (2013)


Subfamily of Centrarchidae in NGW


Following vdLE and our results



Synonym of Rhamphocottidae herein

Following Smith and Busby (2014)



Synonym of Zoarcidae in vdLE

Following Kwan and Kim (2013)


Subfamily of Gadidae in NGW

Subfamily of Lotidae in vdLE

Formerly a subfamily of Lotidae; raised to family level in v3



Subfamily of Balitoridae in vdLE

Following Kottelat (2012)


Subfamily of Kyphosidae in NGW

Subfamily of Kyphosidae in vdLE

Following our results and Carpenter (2001)



Junior synonym of Oxudercidae



Spelled Hapalogeniidae in NGW


See vdLE and Sanciangco et al. (2016)


Subfamily of Percophidae in NGW

Subfamily of Percophidae in vdLE

Following Thacker et al. (2015)



Synonym of Agonidae herein

Following Smith and Busby (2014)


Subfamily of Characidae in NGW


Following Arcila et al. (2017) and vdLE



Subfamily of Cottidae in vdLE

Following Smith and Busby (2014)



Synonym of Gobiidae herein

Following Thacker (2009) and Thacker et al. (2015)


Synonym of Siluridae in NGW


Following vdLE and Britz et al. (2014)


Synonym of Centropomidae herein

Synonym of Centropomidae herein

Following Greenwood (1976) and Li et al. (2011)


Omitted by NGW; listed in erratum





Synomym of Alepocephalidae herein

Following Lavoué et al. (2008) and Paulsen et al. (2009)



Tribe of Paralepididae in vdLE

Following Ghedotti et al. (2014) and Nelson et al. (2016)


Subfamily of Gadidae in NGW


Following Roa-Varón and Ortí (2009)


Synonym (subfamily) of Centriscidae herein


Following vdLE


Synonym (subfamily) of Macrouridae herein


Following Roa-Varón and Ortí (2009)



Synonym of Gobiidae herein

Following Thacker (2009) and Thacker et al. (2015)



Subfamily of Eleotridae in vdLE

Following Thacker (2009) and Thacker et al. (2015)


Subfamily of Scorpaenidae in NGW


Following vdLE


Tribe of Serranidae in NGW

Synonym of Serranidae in vdLE

Following Smith and Craig (2007)


Subfamily of Atherinopsidae herein

Subfamily of Atherinopsidae herein

Following Campanella et al. (2015)


Synonym of Labridae herein

Synonym of Labridae herein

Lumped due to phylogenetic nestedness



Synomym of Bagridae herein

Sullivan et al. (2006)


Synonym of Alepisauridae in NGW


Following Davis (2010)


Omitted by NGW; valid in NEL


Following vdLE



Subfamily of Gobiidae in vdLE

Following Thacker (2009) and Thacker et al. (2015)


Synonym of Bembridae in NGW


Following vdLE and Imamura (1996)



Subfamily of Pleuronectidae in vdLE

Following Chapleau (1993) and Munroe (2005)


Omitted by NGW; subfamily of Kyphosidae in NEL


Following vdLE


Synonym of Percichthyidae herein

Synonym of Percichthyidae herein

Lumped due to phylogenetic nestedness


Not recognized by NGW; Perryena listed under Congiopodidae


Following Honma et al. (2013)



Synomym of Kneriidae herein

Following Davis et al. (2013)


Subfamily of Gadidae in NGW


Following Roa-Varón and Ortí (2009)


Subfamily of Scorpaenidae in NGW


Following Imamura (2004) and vdLE



Subfamily of Pleuronectidae in vdLE

Following Chapleau (1993) and Munroe (2005)


Omitted by NGW; listed in erratum




Synonym (subfamily) of Retropinnidae herein


Following vdLE


Subfamily of Melanotaeniidae in NGW


Following Campanella et al. (2015) and vdLE


Synonym of Gadidae herein


Following Roa-Varón and Ortí (2009)



Subfamily of Pleuronectidae in vdLE

Following Chapleau (1993) and Munroe (2005)


Synonym of Labridae herein

Synonym of Labridae herein

Lumped due to phylogenetic nestedness



Synonym of Gobiidae herein

Following Thacker (2009) and Thacker et al. (2015)


Synonym of Belonidae herein

Synonym of Belonidae herein

Lumped due to phylogenetic nestedness (e.g., Lovejoy et al., 2004)



Subfamily of Cottidae in vdLE

Following Smith and Busby (2014)


Subfamily of Scorpaenidae in NGW


Following vdLE


Subfamily of Scorpaenidae in NGW


Following vdLE



Subfamily of Percichthyidae in vdLE

Following Li et al. (2010)



Subfamily of Merlucciidae in vdLE

Following Roa-Varón and Ortí (2009)



Synonym of Paralepididae in vdLE

Following Davis (2010)



Synomym of Clupeidae herein

Following Lavoué et al. (2013)


Omitted by NGW; valid in NEL


Following vdLE


Subfamily of Scorpaenidae in NGW


Following vdLE


Subfamily of Melanotaeniidae in NGW


Following Campanella et al. (2015) and vdLE


Subfamily of Scorpaenidae in NGW


Following vdLE


Synonym (subfamily) of Macrouridae herein


Following Roa-Varón and Ortí (2009)


Synonym of Iguanodectinae, a subfamily of Characidae in NGW


Following Arcila et al. (2017) and vdLE



Synonym of Eleotridae herein

Following Thacker (2009) and Thacker et al. (2015)


Synonym of Congiopodidae in NGW


Following vdLE



Subfamily of Hexagrammidae in vdLE

Following Smith and Busby (2014)

NEL: Nelson (2006)

NGW: Nelson et al. (2016)

vdLE: van der Laan et al. (2014)


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List of all changes from version 3 (31 July 2014) to version 4 (current). Asterisk (*) denotes 26 changes that were made in accordance with Nelson et al. (2016).

  • Supercohort Elopocephala is no longer classified.
  • Derichthyidae now includes the former Colocongridae*.
  • Osteoglossocephala is now named Osteoglossomorpha*.
  • Clupeidae now includes the round herrings (subfamily Dussumieriinae)*.
  • Alepocephalidae now includes the former Bathylaconidae and Leptochilichthyidae.
  • Suborders in Gonorynchiformes are no longer recognized*.
  • Three otophysan superorders (Cypriniphysae, Characiphysae and Siluriphysae) are now recognized*.
  • “Loricaroidei” has been corrected to “Loricarioidei” (previously misspelled)*.
  • The siluriform families Ailidae, Auchenoglanididae, and Ritidae are now recognized*. 
  • Kryptoglanidae is now recognized.
  • Suborders in Osmeriformes are now classified*.
  • Retropinnidae now includes the former Prototroctidae.
  • Suborders in Stomiatiformes are now classified*.
  • Gonostomatidae now includes the former Diplophidae*.
  • Aulopa is now named Cyclosquamata*.
  • The family Lestidiidae is now recognized*.
  • Endings for the rank “division” have been changed to "-pterygii."
  • Lampridiformes and Lamprididae are now named Lampriformes and Lampridae, respectively*.
  • Zeariae has been corrected to Zeiariae (previously misspelled).
  • Suborders in Zeiformes are now classified*.
  • Zenionidae is now named Zeniontidae*.
  • Euacanthomorphacea is now named Acanthopterygii*.
  • Beryciformes sensu lato is now split into Beryciformes sensu stricto (including suborders Berycoidei and Stephanoberycoidei) and Trachichthyifomes*.
  • Ophidiidae now includes the former Carapidae.
  • Dinematichthyidae is now recognized.
  • Aphyonidae is no longer validated.
  • Pegasidae is now listed as suborder-level incertae sedis in Syngnathiformes (previously in Syngnathoidei).
  • Classification of suborders in Gobiiformes has been modified: Trichonotus is placed in its own suborder (Trichonotoidei); Odontobutoidei and Eleotroidei are now synonyms of Gobioidei.
  • Gobiidae now includes the former Microdesmidae, Kraemeriidae, Ptereleotridae, and Schindleriidae.
  • Butidae, Oxudercidae (= Gobionellidae) and Milyeringidae are now validated.
  • Gobionellidae is now a junior synonym of Oxudercidae*.
  • Cichliformes is expanded herein to include Pholidichthyidae (formerly Pholidichthyiformes)*.
  • Rivulidae is now listed as “Rivulidae.”
  • “Pantanodontidae” is now listed as potential new family 
  • Suborders in Atheriniformes are now classified.
  • Gobiesocidae is now placed in Gobiesociformes (formerly Blenniiformes)*.
  • Gerreiformes is now classified.
  • Labridae now includes the former Odacidae.
  • Centrogenyidae is listed again as order-level incertae sedis in Eupercaria (removed from Uranoscopiformes due to unstable placement).
  • Hapalogenyidae is now listed under Lobotiformes.
  • Sparidae now includes the former Centracanthidae*.
  • Priacanthiformes is now classified.
  • Lujaniformes is now classified.
  • Caproiformes is now classified*.
  • Bathyclupeidae, Champsodontidae and Symphysanodontidae are now listed under Pempheriformes.
  • Pempheriformes “percophids” are now listed as “Percophidae.”
  • Hemerocoetidae is now validated.
  • “Cirrhitioidei” is now corrected to “Cirrhitoidei” (previously misspelled).
  • Dichistiidae is now listed under Terapontoidei.
  • Nine families now included in Scorpaenoidei were listed under Perciformes in previous versions of this classification*.
  • Bembroidei is now named Platycephaloidei and its circumscription is expanded*.
  • Bembropoidei is newly classified.
  • Eulophiidae is now recognized*.
  • Rhamphocottidae now includes the former Ereuniidae*.
  • Family membership for genera in Agonidae, Cottidae and Psychrolutidae is now modified.

List of all changes from version 2 (27 November 2013) to version 3 (31 July 2014)

  • 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. 
  • A new eupercarian order, Chaetodontiformes, is now recognized for the families Chaetodontidae and Leiognathidae. Although support for this clade is weak (50%) in our global 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)).
  • 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.
  • Family Sundasalangidae no longer recognized because Sundasalanx is nested within Clupeidae --following Lavoué et al. (2013) 
  • Family Olyridae no longer recognized. Olyra is included in Bagridae according to Sullivan et al. (2006). 
  • Scoloplacidae is removed from “not examined” in Siluroidei and listed as “not examined” in Loricaroidei.
  • Stathmonotus is now listed within Labrisomidae, following Lin and Hastings (2013). This change renders Chaenopsidae (sensu Lin and Hastings (2013)) monophyletic. 
  • Microdesmus is now listed within Gobiidae, following Thacker, (2009) so Microdesmidae is no longer recognized. 
  • Family Caesionidae (previously listed as insertae sedis in Eupercaria) no longer recognized as valid. It is now listed as synonym of Lutjanidae based on Johnson (1993), Miller and Cribb (2007), and others.
  • Family Scaridae is now included in Labridae, rendering Labridae monophyletic.
  • Family Achiropsettidae no longer recognized. It is considered a synonym of Rhombosoleidae, based on the results presented by Betancur-R. et al. (2013b), rendering Rhombosoleidae monophyletic.
  • Congrogadinae (sensu Godkin and Winterbottom 1985) is now recognized as family Congrogadidae. As a consequence, Pseudochromidae is now monophyletic.
  • 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. 1984Shinohara 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.

List of all changes from version 1 (18 April 2013) to version 2 (27 November 2013)

     Changes based on new taxa examined:
  • Niphonidae: newly validated family, following Smith and Craig (2007); placed in the suborder Percoidei in Perciformes.
  • Phractolaemidae: now examined but no longer recognized as a family; now listed as subfamily of Kneriidae following Davis et al. (2013) .
    • Pegasidae: removed from “not examined” in Callionymoidei to examined in Syngnathoidei. 
    • Zenionidae: now examined; same placement. 
    • Carapidae: now examined; same placement. 
    • Muraenolepididae: now examined; same placement. 
    • Callanthiidae: now examined; listed as incertae sedis in Percomorpharia. 
    • Cepolidae: now examined; listed as incertae sedis in Percomorpharia. 
    • Badidae: now examined; same placement. 
    • Hoplichthyidae: now examined; listed as incertae sedis in Perciformes. 
    • Pentacerotidae: now examined; listed in the newly circumscribed Pempheriformes. 
    • Banjosidae: now examined; listed in the newly circumscribed Pempheriformes. 
    • Ostracoberycidae: now examined; listed in the newly circumscribed Pempheriformes. 
    • Stephanoberycidae: now examined; same placement. 
    • Champsodontidae: now examined; listed as incertae sedis in Percomorpharia. 
    • Aphyonidae: now examined; same placement. 
    • Bembridae: now examined in Perciformes; placed in the newly classified suborder Bembroidei. 
    • Parabembridae: now examined in Perciformes; placed in the newly classified suborder Bembroidei. 
    • Cheimarrichthyidae: now examined; same placement. 
    • Perciliidae: now examined; listed as incertae sedis in Percomorpharia (note that validation of this family results in the paraphyly of Percichthyidae).
    • Pristolepididae: now examined, same placement.
    • Trichodontidae: now examined in Perciformes, infraorder Cottales. 
    • Triodontidae: now examined; placed in the newly classified suborder Triodontoidei. 
    • Anomalopidae: now examined; same placement. 
    • Datnioididae: now examined, placed in the newly classified order Lobotiformes (with Lobotidae). 
    • Phallostethidae: now examined; same placement. 

         Other Changes:

    • Creediidae is removed from Syngnathiformes and placed in Pempheriformes (Percomorpharia). This is the major topological change in the new tree. The previous version had only four gene sequences for Limnichthys (two of which were likely contaminated) and this species is now represented in eight markers. The new placement is consistent with that obtained by Near et al. (2013) .
    • The order Pempheriformes has a broader circumscription, including the families Acropomatidae, Banjosidae, Creediidae, Epigonidae, Glaucosomatidae, Howellidae, Lateolabracidae, Leptoscopidae, Ostracoberycidae, Pempheridae, Pentacerotidae, Percophidae, and Polyprionidae. 
    • Bembrops is now placed in Bembropidae, following Near et al. (2013). Percophidae, as traditionally recognized (including Bembropinae), is polyphyletic in the new tree (as well as in Near et al., 2013) , with Acanthaphristis and Bembrops falling in different places. 
    • Girella is now recognized in its own family, Girellidae, following Carpenter (2001) . Inclusion of Girella in Kyphosidae renders this family non-monophyletic. 
    • The families Girellidae, Kuhliidae, Kyphosidae, Oplegnathidae, and Terapontidae are classified in the new order Terapontiformes in Percomorpharia. 
    • Fistulariidae is removed from “possibly included” in Syngnathoidei and listed as incertae sedis in Syngnathiformes. 
    • Leptoscopidae is removed from not examined” in Syngnathiformes (Syngnathimorpharia) and listed in the newly circumscribed Pempheriformes (Percomorpharia). 
    • Pelagimorpharia was adopted to replace Scombrimorpharia, following Miya et al. (2013)Pelagia was not adopted because it is already in use for a genus of scyphozoan jellyfish (Fam Pelagiidae). 
    • Six families traditionally placed in “Perciformes”, and previously listed as incertae sedis in Percomorphaceae, are provisionally listed in Percomorpharia: Bathyclupeidae, Dichistiidae, Hapalogenyidae, Parascorpididae, Symphysanodontidae, Trichonotidae.
    • The superorder “Cyprinae” was changed to “Cypriniphysae” following Nelson (2006).
    •  The monophyly of both Ceratioidei and Ceratiidae was challenged in version 1 due to misplacement of Cryptosaras. This is no longer the case in the phylogeny supporting version 2, where Cryptosaras and Ceratias are resolved as sister taxa with 98% bootstrap support.

    Ricardo Betancur,
    Jul 31, 2016, 8:09 AM
    Ricardo Betancur,
    Jul 31, 2016, 8:09 AM
    Ricardo Betancur,
    Jul 30, 2016, 11:59 AM
    Ricardo Betancur,
    Jul 31, 2016, 8:10 AM