Version Date: 27 November 2013 -- this version has been superseded by version 3 on 31 July 2014
Citing this classification:
Betancur-R, R., E. Wiley, M. Miya, G. Lecointre, N. Bailly, and G. Ortí. 2013. New and Revised Classification of Bony Fishes Version 2 (http://www.deepfin.org/Classification_v2.htm).
Betancur-R., R., R.E. Broughton, E.O. Wiley, K. Carpenter, J.A. Lopez, C. Li, N.I. Holcroft, D. Arcila, M. Sanciangco, J. Cureton, F. Zhang, T. Buser, M. Campbell, T. Rowley, J.A. Ballesteros, G. Lu, T. Grande, G. Arratia & G. Ortí. 2013. The tree of life and a new classification of bony fishes. PLoS Currents Tree of Life. 2013 Apr 18.
This classification is an update of version 1 (18 April 2013) published by Betancur-R. et al. (2013a) . It is based on new analyses of a molecular data set with 1591 taxa that adds 165 new taxa and ca. 25 families not examined in version 1. The new data set combines evidence published by Betancur-R. et al. (2013a) and Near et al. (2013). Methods used for phylogenetic inference were as described by Betancur-R. et al. (2013a), based on maximum likelihood analyses with RAxML using a 24-partition scheme and 1000 bootstrap replicates to assess clade support. The complete phylogenetic tree (pdf, unpublished results), a pretty figure with all the major groups, and a spreadsheet with the classification scheme can be downloaded at the bottom of this page. Families in the classification are linked to FishBase family pages (Froese and Pauly (2013); except for the differences noted in the spreadsheet).A major change in version 2 is that the nine well-supported percomorph clades are now recognized as a separate series to facilitate communication. These are: Ophidiimorpharia, Batrachoidimorpharia, Gobiomorpharia, Syngnathimorpharia, Pelagimorpharia (Scombriformes in version1), Anabantomorpharia, Carangimorpharia, Ovalentaria, and Percomorpharia (Fig. 1). A complete list of changes from version 1 can be found here.
Figure 1. Phylogeny of the nine major percomorph (Percomorphaceae) clades (from Betancur-R. et al. (2013a) and new results). Each major clade is now recognized as a separate series. Values in parentheses indicate total number of orders, total number of families, and the number of families included in each series that cannot be placed in any order at this time (insertae sedis families).
This classification (version 2), building on Wiley and Johnson (2010) and Betancur-R. et al. (2013a), 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 previous molecular studies (several examples discussed below). Order-level or supraordinal taxa have been erected (listed as new) or resurrected on the basis of well-supported clades only (>90% bootstrap values). Current taxon names supported by previous molecular or morphological studies have been retained if congruent with our results, even if bootstrap support is low (e.g., Osteoglossocephalai sensu Arratia (1999) with only 38% bootstrap). In some cases, ordinal or subordinal taxa that were not monophyletic in our analysis are also validated, as long as the incongruence is not supported by strong bootstrap values. Examples include the suborder Blennioidei (not monophyletic here but monophyletic in Wainwright et al., 2012) and the order Pleuronectiformes (not monophyletic here but monophyletic in Betancur-R. et al., 2013b).
A total of 68 orders are classified, of which two are new for version 2 (Lobotiformes and Terapontiformes). Three orders were new in the version 1 (Holocentriformes, Istiophoriformes, and Pempheriformes). The ordinal status of several percomorph families examined (as well as many others unexamined) belonging to the Series Carangimorpharia, Ovalentaria, and Percomorpharia remains uncertain (incertae sedis) due to poor phylogenetic resolution. We therefore list these families as incertae sedis within each of these groups (Carangimorpharia, Ovalentaria, and Percomorpharia) awaiting new phylogenetic evidence to clarify their ordinal status.
Family names for bony fishes are based on Eschmeyer and Fong (2013) and van der Laan et al. (2013), with minor modifications. Van der Laan et al. (2013) and Eschmeyer (2013) should be consulted for authorship of family names and Wiley and Johnson (2010) for authorship of ordinal and subordinal names. Our list is not intended as a comprehensive revision of valid family names. Instead, it is simply an adaptation of their lists based on published studies that we know validate or synonymize family groups using explicit phylogenetic evidence. Following this criterion, we do not recognize the family status of Phractolaemidae (synonym of Kneriidae; Davis et al., 2013), Anotopteridae, Omosudidae (synonym of Alepisauridae; Davis, 2010), or Latidae (synonym of Centropomidae; Greenwood, 1976; Li et al., 2011). Also, we recognize the following families, listed in Eschmeyer and Fong (2013) and van der Laan et al. (2013) as synonyms or subfamilies of other families: Botiidae (Chen et al., 2009), Diplophidae (Nelson, 2006 apparently ommited by ; Eschmeyer and Fong, 2013), Horabagridae (Sullivan et al., 2006), Niphonidae (Smith and Craig, 2007), Sinipercidae (Li et al., 2010), Steindachneriidae (Roa-Varon and Orti, 2009), Sudidae (Davis, 2010), Bembropidae (Near et al., 2013), and the pleuronectiform Paralichthodidae, Poecilopsettidae, and Rhombosoleidae (Chapleau, 1993; Munroe, 2005).
A total of 504 families of bony fishes are now recognized (excluding tetrapods), of which 396 (78.6%) were examined. Among the examined families, 41 (8.1 %) were rendered non-monophyletic in our analysis (indicated below). For each order/suborder we list all families examined as well as the unexamined families whose taxonomic affinity is expected on the basis of traditional taxonomy or phylogenetic evidence. The list of 108 unexamined families can be easily obtained from the spreadsheet that also contains the complete classification (bottom of this page), and is intended as a resource to help fish systematists direct future sequencing efforts.
The new classification scheme presented here should still be considered work in progress (version 2), 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 email@example.com. For the most updated version always visit DeepFin.
Classification for Bony Fishes (version 2)
Megaclass Osteichthyes (=Euteleostomi, =Euosteichthyes)
Superclass Actinopterygii (100%)
Class Cladistia (100%)
Class Actinopteri (100%)
Subclass Chondrostei, (100%)
Subclass Neopterygii (100%)
Infraclass Holostei (100%)
Order Lepisosteiformes (100%)
Infraclass Teleostei (100%)
Megacohort Elopocephalai sensu Arratia (1999) (100%)
Supercohort Elopocephala (100%)
Cohort Elopomorpha (100%)
Order Elopiformes (100%)
Order Albuliformes (100%)
Order Notacanthiformes (100%)
Order Anguilliformes (100%)
Comment: Suborders recognized in Wiley and Johnson (2010) based on previous work cited therein are significantly incongruent with the clades obtained in this analysis; thus, no subordinal classification is proposed.
Megacohort Osteoglossocephalai sensu Arratia (1999) (38%)
Supercohort Osteoglossocephala sensu Arratia (1999) (99%)
Order Hiodontiformes (100%)
Order Osteoglossiformes (100%)
Supercohort Clupeocephala sensu Arratia (2010) (100%)
Cohort Otomorpha (=Otocephala, Ostarioclupeomorpha) (100%)
Subcohort Clupei (87%)
Suborder Clupeoidei (100%)
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.
Subcohort Alepocephali (100%)
Alepocephalidae (not monophyletic)
Not examined: Leptochilichthyidae.
Subcohort Ostariophysi (100%)
Section Anotophysa (97%)
Suborder Knerioidei (100%)
Section Otophysa (100%)
Superorder Cypriniphysae (100%)
Superorder Characiphysae (100%)
Order Gymnotiformes (100%)
Suborder Gymnotoidei (not monophyletic)
Gymnotidae (not monophyletic)
Suborder Sternopygoidei (not monophyletic)
Sternopygidae (not monophyletic)
Not examined: Hypopomidae.
Comment: Although not monophyletic here, the monophyly gymnotiform suborders is corroborated by Albert and Crampton (2005).
Order Characiformes (100%)
Suborder Citharinoidei (not monophyletic)
Suborder Characoidei (not monophyletic)
Comment: Although not monophyletic in this analysis, the monophyly of characiform suborders has been corroborated by other molecular studies (Calcagnotto et al., 2005).
Order Siluriformes (100%)
Suborder Loricaroidei (93%)
Suborder Siluroidei (100%)
Supercohort Clupeocephala (cont.)
Cohort Euteleosteomorpha (100%)
Subcohort Protacanthopterygii sedis mutabilis (37%)
Order Galaxiiformes (100%)
Order Argentiniformes (100%)
Order Salmoniformes (100%)
Order Esociformes (100%)
Subcohort Stomiatii (73%)
Order Stomiatiformes (=Stomiiformes) (100%)
Phosichthyidae (not monophyletic)
Order Osmeriformes (100%)
Subcohort Neoteleostei (100%)
Infracohort Ateleopodia (100%)
Infracohort Eurypterygia (96%)
Section Aulopa (100%)
Suborder Aulopoidei (not monophyletic)
Synodontidae (not monophyletic)
Suborder Alepisauroidei (not monophyletic)
Chlorophthalmidae (not monophyletic)
Ipnopidae (not monophyletic)
Paralepididae (not monophyletic)
Scopelarchidae (not monophyletic)
Section Ctenosquamata (97%)
Subsection Myctophata (100%)
Subsection Acanthomorphata (97%)
Division Lampridacea (100%)
Division Paracanthomorphacea sensu Grande et al. (2013) (93%)
Series Percopsaria (100%)
Series Zeiogadaria (=Zeiogadiformes sensu Li et al., 2009) (98%)
Subseries Zeariae (100%)
Subseries Gadariae (100%)
Order Stylephoriformes (sensu Miya et al., 2007)
Order Gadiformes (100%)
Macrouridae (not monophyletic)
Suborder Gadoidei (not monophyletic)
Lotidae (not monophyletic)
Comment: The subordinal classification follows Roa-Varón and Orti (2009): fig. 6.
Division Polymixiacea (100%)
Division Euacanthomorphacea sensu Johnson and Patterson (1993) (99%)
Subdivision Berycimorphaceae (87%)
Order Beryciformes (similar to Trachichthyiformes sensu Moore, 1993)
Trachichthyidae (not monophyletic)
Subdivision Holocentrimorphaceae (100%)
Comment: Moore (1993) and Stiassny and Moore (1992) provide morphological evidence supporting a sister-group relationship between holocentrids and percomorphs, which further guarantees placement of this family in its own order.
Subdivision Percomorphaceae (cont.)
Series Ophidiimorpharia (100%)
Carapidae (not monophyletic)
Bythitidae (not monophyletic)
Not examined: Parabrotulidae
Series Batrachoidimorpharia (100%)
Comment: In addition to the well-supported molecular circumscription of this group, kurtids, apogonids and gobioids are characterized by the presence of sensory papillae rows on the head and body (Thacker, 2009).
Order Kurtiformes (98%)
Comment: Johnson (1993) noted that the configuration of the dorsal gill-arch elements may be homologous in Kurtus and apogonids.
Order Gobiiformes (100%)
Suborder Odontobutoidei (100%)
Suborder Eleotroidei (97%)
Suborder Gobioidei (100%)
Gobiidae (not monophyletic)
Series Syngnathimorpharia, new (96%)
Suborder-level incertae sedis in Syngnathiformes
Suborder Syngnathoidei (92%)
Not examined: Solenostomidae (assumed affinity with Syngnathidae)
Suborder Dactylopteroidei (100%)
Suborder Callionymoidei (100%)
Not examined: Draconettidae (assumed affinity with Callionymidae).
Suborder Mulloidei (92%)
Gempylidae (not monophyletic)
Scombridae (not monophyletic)
Comment: interfamilial resolution in Scombriformes is tenuous; circumscription of scombriform families into suborders (e.g., Scombroidei, Stromateoidei, Icostoidei) or new orders requires further work.
Series Anabantomorpharia (=Anabantiformes sensu Li et al., 2009) (99%)
Order Synbranchiformes (98%)
Not examined: Chaudhuriidae.
Order Anabantiformes (=Labyrinthici) (100%)
Suborder Anabantoidei (95%)
Suborder Channoidei (85%)
Series Carangimorpharia (=Carangimorpha sensu Li et al. (2009)) (100%)
Order-level incertae sedis in Carangimorpharia
Centropomidae (sensu Greenwood)
Order Istiophoriformes (100%)
Order Carangiformes sedis mutabilis (not monophyletic)
Comment: Monophyly of Carangiformes is not significantly rejected by the data (see Betancur-R. et al., 2013b).
Order Pleuronectiformes sedis mutabilis (not monophyletic)
Suborder Psettodoidei (100%)
Suborder Pleuronectoidei (100%)
Paralichthyidae (not monophyletic)
Rhombosoleidae (not monophyletic)
Not examined: Paralichthodidae.
Comment: Although Psettodidae is not recovered as the sister group of pleuronectoids in the present analysis, the order was resolved as monophyletic by a recent study (Betancur-R. et al., 2013b).
Order-level incertae sedis in Ovalentaria
Grammatidae (not monophyletic)
Pseudochromidae (not monophyletic)
Superorder Cichlomorphae (93%)
Superoder Atherinomorphae (100%)
Order Atheriniformes (100%)
Order Beloniformes (32%)
Suborder Exocoetoidei (100%)
Belonidae (not monophyletic)
Hemiramphidae (not monophyletic)
Zenarchopteridae (not monophyletic)
Order Cyprinodontiformes (57%)
Suborder Cyprinodontoidei (100%)
Superorder Mugilomorphae (100%)
Superorder Blenniimorphae (80%)
Order Blenniiformes (100%) sensu Li et al. (2009)
Suborder Blennioidei (not monophyletic; = Blenniiformes sensu Lin and Hastings (2013))
Chaenopsidae (not monophyletic)
Labrisomidae (not monophyletic)
Comment: While blennioids are not monophyletic in the megatree, we note that preliminary analyses resulted in the reciprocal monophyly of gobiesocoids and blennioids, which is congruent with molecular (Wainwright et al., 2012; Lin and Hastings, 2013) and morphological (Springer and Orrell, 2004) evidence. Monophyly of gobiesocoids and blennioids (as separate orders/suborders) is further supported by both morphological (Wiley and Johnson, 2010) and molecular evidence.
Series Percomorpharia (99%)
Order-level incertae sedis in Percomorpharia
Lutjanidae (not monophyletic)
Percichthyidae (not monophyletic)
Not examined: Dinolestidae, Dinopercidae (see Smith and Craig, 2007). Six families traditionally placed in “Perciformes” are also provisionally listed here are: Bathyclupeidae, Dichistiidae, Hapalogenyidae, Parascorpididae, Symphysanodontidae, Trichonotidae; these are not placed in the recently circumscribed Perciformes given the long history of phylogenetic indistinctiveness between Percoidei, Perciformes, and Percomorpha (e.g., Smith and Craig, 2007).
Order Uranoscopiformes (=Paratrachinoidei sensu Li et al. (2009)) (95%)
Order Labriformes sensu stricto (100%)
Labridae (not monophyletic)
Order Lobotiformes (100%)
Order Ephippiformes (100%)
Comment: Greenwood et al. (1966) hypothesized a close affinity between Drepane and ephippids.
Possibly included (examined): Nemipteridae
Comment: Akazaki (1962) proposed that Lethrinidae, Sparidae, and Nemipteridae were closely related based on specializations of the suspensorium and other features (Johnson, 1993). Johnson (1981) supported the monophyly of Akazaki's spariforms with the addition of Centracanthidae.
Order 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%)
Order Tetraodontiformes (100%). This order is the sister group of Lophiiformes (55% bootstrap); also supported by anatomical evidence (Chanet et al. (2013), larval characters (Baldwin (2013)), and previous molecular studies (e.g. Dettaï and Lecointre (2008), Miya et al. (2003), Miya et al., (2010)).
Suborder Triacanthodoidei (100%)
Suborder Tetraodontoidei (100%)
Suborder Moloidei (100%)
Suborder Balistoidei (100%)
Suborder Ostracioidei (100%)
Suborder Triodontoidei, new
Comment: This subordinal classification differs from that proposed by Santini and Tyler (2003).
Order Acanthuriformes, restricted circumscription (see also Holcroft and Wiley, 2008) (100%)
Order Terapontiformes, new (95%)
Order Pempheriformes sedis mutabilis, new circumscription (44%)
Acropomatidae (not monophyletic)
Polyprionidae (not monophyletic)
Comment: Tominaga (1986) suggested that features of the cranium and swimbladder may be homologous in Pempheris and Glaucosoma. Although support for Pempheriformes is only 44%, this clade is often recovered in different analyses.
Order Centrarchiformes (100%)
Not examined (12 families traditionally placed in Scorpaeniformes): Apistidae, Aploactinidae, Congiopodidae, Cottocomephoridae, Eschmeyeridae, Gnathanacanthidae, Neosebastidae, Pataecidae, Perryenidae, Plectrogeniidae, Zanclorhynchidae.
Suborder-level incertae sedis in Perciformes
Suborder Serranoidei sedis mutabilis (19%)
Suborder Percoidei, restricted circumscription (99%)
Not examined: Trachinidae.
Comment: Lautredou et al. (2013) using seven nuclear markers obtained a clade uniting Percidae and Trachinidae with full support.
Suborder Notothenioidei (100%)
Nototheniidae (not monophyletic)
Bathydraconidae (not monophyletic)
Suborder Scorpaenoidei (83%)
Scorpaenidae (not monophyletic)
Suborder Bembroidei (96%)
Suborder Triglioidei sensu Jordan (1923) (100%)
Suborder Cottioidei (=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 most probably sister-groups; they are have been grouped as Zoarciformes by Li et al. (2009)
Infraorder Gasterosteales (100%)
Infraorder Zoarcales (100%)
Bathymasteridae (not monophyletic)
Stichaeidae (not monophyletic)
Infraorder Cottales (96%)
Agonidae (not monophyletic)
Cottidae (not monophyletic)
Hexagrammidae (not monophyletic)
Superclass Sarcopterygii (96%)
Class Coelacanthimorpha (=Actinistia)
Class Dipnotetrapodomorpha sedis mutabilis (65%)
Subclass Dipnomorpha (100%)
Suborder Lepidosirenoidei (100%)
Subclass Tetrapodomorpha (100%)
Changes based on new taxa examined:
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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, and G. Ortí. 2013a. The tree of life and a new classification of bony fishes. PLoS Currents Tree of Life. 2013 Apr 18.
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Burridge, C. P., and A. J. Smolenski. 2004. Molecular phylogeny of the Cheilodactylidae and Latridae (Perciformes: Cirrhitoidea) with notes on taxonomy and biogeography. Molecular Phylogenetics and Evolution. 30:118-127.
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Carpenter, K. 2001. Girellidae, Scorpididae, Microcanthidae, p. 2791-3379. In: The Living Marine Resources of the Western Central Pacific. FAO Species Identification Guide for Fisheries Purposes. Vol. 5. K. Carpenter and V. H. Niem (eds.). FAO, Rome.
Chanet, B., Guintard, C., Betti, E., Gallut, C., Dettaï, A. and G. Lecointre. 2013. Evidence for a close phylogenetic Relationship between the teleost orders Tetraodontiformes and Lophiiformes based on an analysis of soft anatomy. Cybium 37:179-198.
Chen, W. J., V. Lheknim, and R. L. Mayden. 2009. Molecular phylogeny of the Cobitoidea (Teleostei: Cypriniformes) revisited: position of enigmatic loach Ellopostoma resolved with six nuclear genes. Journal of Fish Biology. 75:2197-2208.
Davis, A. M., G. Arratia, and T. M. Kaiser. 2013. The first fossil shellear and its implications for the evolution and divergence of the Kneriidae (Teleostei: Gonorynchiformes), p. 325-362. In: Mesozoic Fishes 5 - Global Diversity and Evolution. G. Arratia, H.-P. Schultze, and M. V. H. Wilson (eds.). Verlag F. Pfeil, Muenchen.
Davis, M. P. 2010. Evolutionary relationships of the Aulopiformes (Euteleostei: Cyclosquamata): a molecular and total evidence approach, p. 317-336. In: Origin and Phylogenetic Interrelationships of Teleosts. J. S. Nelson, H. P. Schultze, and M. V. H. Wilson (eds.). Verlag Dr. Friedrich Pfeil, München, Germany.
Dettaï, A., G. Lecointre. 2004. New insights into the organization and evolution of vertebrate IRBP genes and utility of IRBP gene sequences for the phylogenetic study of the Acanthomorpha (Actinopterygii : Teleostei). Molecular Phylogenetics and Evolution 48:258-269.
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Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers. 1966. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bulletin of the American Museum of Natural History. 131.
Lautredou, A.-C., H. Motomura, C. Gallut, C. Ozouf-Costaz, C. Cruaud, G. Lecointre, and A. Dettai. 2013. Multi-scale exploration of the relationships among Serraniformes (Acanthomorpha, Teleostei) using new nuclear markers. Molecular Phylogenetics and Evolution. 67:140-155.
Lavoue, S., M. Miya, P. Musikasinthorn, W.-J. Chen, M. Nishida. 2013.Mitogenomic evidence for an Indo-West Pacific origin of the Clupeoidei (Teleostei: Clupeiformes). PLoS ONE 8(2): e56485. doi:10.1371/journal.pone.0056485.
Li, C., R. Betancur-R., W. L. Smith, and G. Orti. 2011. Monophyly and interrelationships of Snook and Barramundi (Centropomidae sensu Greenwood) and five new markers for fish phylogenetics. Molecular Phylogenetics and Evolution. 60:463-71.
Miya, M., M. Friedman, T. P. Satoh, H. Takeshima, T. Sado, W. Iwasaki, Y. Yamanoue, M. Nakatani, K. Mabuchi, J. G. Inoue, J. Y. Poulsen, T. Fukunaga, Y. Sato, and M. Nishida. 2013. Evolutionary origin of the scombridae (tunas and mackerels): members of a paleogene adaptive radiation with 14 other pelagic fish families. PLoS ONE. 8:e73535.
Miya, M., N. I. Holcroft, T. P. Satoh, M. Yamaguchi, M. Nishida, and E. O. Wiley. 2007. Mitochondrial genome and a nuclear gene indicate a novel phylogenetic position of deep-sea tube-eye fish (Stylephoridae). Ichthyological Research. 54.
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Miya, M., H. Takeshima, H. Endo, N.B. Ishiguro, J.G. Inoue, T. Mukai, T.P. Satoh, M. Yamaguchi, A. Kawaguchi, K. Mabuchi, S.M.N. Shirai and M. Nishida. 2003. Major patterns of higher teleostean phylogenies: a new perspective based on 100 complete mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 26:121–138.
Near, T. J., A. Dornburg, R. I. Eytan, B. P. Keck, W. L. Smith, K. L. Kuhn, J. A. Moore, S. A. Price, F. T. Burbrink, M. Friedman, and P. C. Wainwright. 2013. Phylogeny and tempo of diversification in the superradiation of spiny-rayed fishes. Proceedings of the National Academy of Sciences 101:12738-21743. doi: 10.1073/pnas.1304661110.
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Springer, V. G., and T. M. Orrell. 2004. Phylogenetic analysis of the families of acanthomorph fishes based on dorsal gill-arch muscles and skeleton. Bulletin of the Biological Society of Washington. 11:237–260.
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