Version Date: XXX 2016
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.
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General comments on the classification
This classification is an update of previous versions (version 1, version 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 2. Van 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 1) belonging 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 tree. Examples 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.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%)
Class Actinopteri (100%)
Subclass Chondrostei (100%)
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%)
Supercohort Clupeocephala sensu Arratia (2010) (100%)
Cohort Otomorpha (= Otocephala, Ostarioclupeomorpha) (92%)
Subcohort Clupei (= Clupeomorpha) (100%)
Order Clupeiformes (100%)
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%)
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%)
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., 2017) provides 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%)
Superorder Characiphysae (= Characiphysi) (100%)
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 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 Siluroidei (100%)
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 Protacanthopterygii sedis mutabilis (100%)
Order Galaxiiformes (94%)
Order Argentiniformes (47%)
Order Salmoniformes (62%)
Order Esociformes (100%)
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%)
Subcohort Neoteleostei (100%)
Infracohort Ateleopodia (= Ateleopodomorpha) (98%)
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).
Suborder Aulopoidei (not monophyletic)
Synodontidae (not monophyletic)
Suborder Alepisauroidei (not monophyletic)
Chlorophthalmidae (not monophyletic)
Ipnopidae (not monophyletic)
Paralepididae (not monophyletic)
Scopelarchidae (not monophyletic)
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%)
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)
Comment: Endings for the rank Division have been changed to "-pterygii" (see comments under Acanthopterygii below).
Series Percopsaria (100%)
Series Zeiogadaria (= Zeiogadiformes sensu Li et al. (2009)) (100%)
Subseries Zeiariae (100%)
Subseries Gadariae (100%)
Order Stylephoriformes (sensu Miya et al. (2007))
Order Gadiformes (100%)
Suborder Macrouroidei (100%)
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)
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).
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 lineages. This 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%)
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 Stephanoberycoidei) and 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%)
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 (cont.)
Series Ophidiaria (100%)
Suborder Ophidioidei (100%)
Suborder Bythitoidei (100%)
Bythitidae (not monophyletic)
Not examined: Parabrotulidae.
Series Batrachoidaria (100%)
Series Syngnatharia (84%)
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%)
Gempylidae (not monophyletic)
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.
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%)
Not examined: Trichonotidae.
Suborder Gobioidei (100%)
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%)
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
Order Istiophoriformes (100%)
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)
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").
Order-level incertae sedis in Ovalentaria
Ambassidae (= Chandidae)
Superorder Cichlomorphae (40%)
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%)
Order Beloniformes (79%)
Suborder Belonoidei (100%) (= Exocoetoidei sensu Nelson et al. (2016))
Hemiramphidae (not monophyletic)
Order Cyprinodontiformes (100%)
Suborder Cyprinodontoidei (100%)
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., 2014; Pohl 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%)
Superorder Blenniimorphae (90%)
Order Gobiesociformes (100%)
Order Blenniiformes (39%)
Chaenopsidae (not monophyletic)
Comments: Circumscription of Blenniiformes follows Lin and Hastings (2013). The reciprocal monophyly of gobiesocoids and blennioids is supported by molecular (Wainwright et al., 2012; Lin 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
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%)
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 Acanthuridae, Luvaridae and Zanclidae. Our results do not support the placement of Emmelichthyidae and Sciaenidae in Acanthuriformes.
Order Lutjaniformes, new circumscription (59%)
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.
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 Callanthiidae, Lobotidae (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%)
Suborder Chaunacoidei (100%)
Suborder Ogcocephaloidei (100%)
Suborder Ceratioidei (100%)
Order Tetraodontiformes (100%). This order is the sister group of Lophiiformes (see comments under Lophiiformes above).
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)
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, Pteropsaron, Acanthaphritis, and Osopsaron and the unsampled Dactylopsaron, Hemerocoetes, and Squamicreedia.”
Order Centrarchiformes (98%)
Suborder Centrarchoidei (93%)
Comment: Inclusion of Enoploside in this suborder differs from the results obtained by Lavoué et al. (2014).
Cheilodactylidae (not monophyletic)
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%)
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).
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).
Not examined: Normanichthyidae.
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%)
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)
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%)
Comment: Previous versions of this classification included Bembridae and Parabembridae in the suborder Bembroidei, which we now expand to also include Hoplichthyidae, Platycephalidae 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)
Infraorder Gasterosteales (similar Gasterosteoidei in other classifications, but excluding Indostomidae) (100%)
Infraorder Zaniolepidoales (= Zaniolepidoidei sensu Smith and Busby (2014))
Infraorder Hexagrammales (100%) (= Hexagrammoidei in previous classifications)
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))
Scorpaenichthyidae (following Smith 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)
Class Dipnotetrapodomorpha (100%)
Superorder Ceratodontae (= Dipnoi)
Suborder Lepidosirenoidei (100%)
Subclass Tetrapodomorpha (100%)
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