Source: http://www.fishphylogeny.org/classification
Timestamp: 2019-04-18 11:18:51+00:00

Document:
Betancur-R, R., E. Wiley, N. Bailly, M. Miya, G. Lecointre, and G. Ortí. 2014. Phylogenetic Classification of Bony Fishes --Version 3 (http://www.deepfin.org/Classification_v3.htm).
Betancur-R., R., R.E. Broughton, E.O. Wiley, K. Carpenter, J.A. Lopez, C. Li, N.I. Holcroft, D. Arcila, M. Sanciangco, J. Cureton, F. Zhang, T. Buser, M. Campbell, T. Rowley, J.A. Ballesteros, G. Lu, T. Grande, G. Arratia & G. Ortí. 2013. The tree of life and a new classification of bony fishes. PLoS Currents Tree of Life. 2013 Apr 18.
Go to the bottom of this page to download: Spreadsheet with complete classification scheme; molecular phylogeny used as basis for classification (from version 2, needs to be updated to reflect the current classification scheme); and figures with summary phylogenetic hypothesis displaying all major groups.
This classification is an update of version 1 (18 April 2013) and version 2 (27 Nov 2013), originally published by Betancur-R. et al. (2013a). This version fixes involuntary errors and omissions (thanks to numerous colleagues who provided feedback!) and incorporates new findings from the recent literature. It is based on the same phylogenetic results used for version 2, from analysis of a molecular data set with 1591 taxa (the data set combines evidence published by Betancur-R. et al. (2013a) and Near et al. (2013)). Families in the classification are linked to FishBase family pages (Froese and Pauly, 2013) except for cases where there are discrepancies in the recognition of valid families, noted and justified in each case in the classification.
Version 3 presents new names for series included in subdivision Percomorphaceae, to avoid confusion between this taxon and series Percomorpharia, and hence disambiguate the meaning of "percomorph". Percomorpharia is now referred to as series Eupercaria, and the other series names also have simpler endings (dropping the term "morph" but keeping a consistent ending with "aria"). Under this scheme, the word "percomorph" refers unambiguously to subdivision Percomorphaceae, retaining its original meaning as implied by Wiley and Johnson (2010), following Johnson and Patterson (1993). The common name "eupercarians" now is available for taxa in the clade containing Perciformes, Labriformes, Tetraodontiformes and others (i.e., the "new bush at the top" sensu Betancur-R. et al. (2013a)). Figure 1 below shows the new names.
A complete list of changes from version 2 can be found here.
General comments on the classification are here.
Figure 1. Phylogeny of the nine major percomorph clades included in Subdivision Percomorphaceae. Each major clade is recognized as a series.
Not examined: Chlopsidae, Colocongridae, Cyematidae, Derichthyidae, Heterenchelyidae, Monognathidae, Moringuidae, Myrocongridae, Nettastomatidae, Protanguillidae, Synaphobranchidae.
Comment: Suborders recognized in Wiley and Johnson (2010) based on previous work cited therein are significantly incongruent with the clades obtained in this analysis; thus, no subordinal classification is proposed.
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.
Comment: the position of alepocephaloids as the sister group to Ostariophyisi also was reported by Lavoué et al. (2008) and by Paulsen et al. (2009).
Not examined: Balitoridae, Barbuccidae, Ellopostomatidae, Psilorhynchidae, Serpenticobitidae, Vaillantellidae.
Comment: Although not monophyletic here, the monophyly of gymnotiform suborders is corroborated by Albert and Crampton (2005).
Not examined: Anostomidae, Curimatidae, Iguanodectidae.
Comment: Although not monophyletic in this analysis, the monophyly of characiform suborders has been corroborated by other molecular studies (Calcagnotto et al., 2005). In other molecular studies, the monophyly of Characiformes was not obtained, since Characoidei was more closely related to Siluriformes than to Citharinoidei (Nakatani et al., 2011; Chen et al., 2014a).
Not examined: Aspredinidae, Austroglanididae, Erethistidae, Lacantuniidae.
Not examined: Bathysauroididae, Bathysauropsidae sensu Davis (2010).
Comment: Anotopteridae is not recognized; aulopiform families listed follow Davis (2010). Although not monophyletic herein, the monophyly of aulopiform suborders listed is supported by Davis (2010).
Not examined: Cyttidae, Grammicolepididae, Oreosomatidae.
Not examined: Bregmacerotidae, Euclichthyidae, Melanonidae.
Comment: The subordinal classification follows Roa-Varón and Ortí (2009): fig. 6.
Not examined: Gibberichthyidae, Hispidoberycidae (expected affinity following Moore (1993)).
Comment: Moore (1993) and Stiassny and Moore (1992) provide morphological evidence supporting a sister-group relationship between holocentrids and percomorphs, which further guarantees placement of this family in its own order.
Subdivision Percomorphaceae (="Percomorpha" sensu Miya et al. (2003) and Miya et al. (2005)).
Comment: Johnson (1993) noted that the configuration of dorsal gill-arch elements may be homologous in Kurtus and apogonids.
Not examined: Kraemeriidae, Rhyacichthyidae, Schindleriidae, Thalasseleotrididae, Xenisthmidae.
Comment: In addition to the well-supported molecular circumscription of Gobiaria, kurtids, apogonids and gobioids are characterized by the presence of sensory papillae rows on the head and body (Thacker, 2009).
Not examined: Draconettidae (assumed affinity with Callionymidae).
Comment: interfamilial resolution in Scombriformes is tenuous; circumscription of scombriform families into suborders (e.g., Scombroidei, Stromateoidei, Icostoidei) or new orders requires further work.
Comment: suborders of Anabantiformes now reflect well-supported monophyletic groups, correcting an error in Version 2 and defining a third suborder (Channoidei) for the family Channidae. Affinities of Channidae with other anabantiform families varies among studies (e.g., Near et al. (2013), Betancur-R. et al. (2013a), and our new results). The new scheme with three subroders is robust to this ambiguity.
Comment: Monophyly of Carangiformes is not significantly rejected by the data (Betancur-R. et al., 2013b).
Not examined: Paralichthodidae (following Chapleau (1993) and Munroe (2005)).
Comment: Although Psettodidae is not recovered as the sister group of pleuronectoids in the present analysis, the order was resolved as monophyletic by recent studies that address this issue specifically (Betancur-R. et al. (2013b), Betancur-R and Ortí (2014)).
Not examined: Anablepidae, Goodeidae, Profundulidae, Valenciidae.
Comment: While blennioids are not monophyletic in our results, we note that preliminary analyses resulted in the reciprocal monophyly of gobiesocoids and blennioids, which is congruent with molecular (Wainwright et al., 2012; Lin and Hastings, 2013) and morphological (Springer and Orrell, 2004) evidence. Monophyly of gobiesocoids and blennioids (as separate orders/suborders) is further supported by both morphological (Wiley and Johnson, 2010) and molecular evidence. Chaenopsidae is monophyletic if Stathmonotus is included in Labrisomidae, following Lin and Hastings (2013).
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)).
Possibly included: Centrogenyidae (bootstrap support for Uranoscopiformes plus Centrogenyidae is only 41%, as opposed to 99% for Uranoscopiformes s.s. but placement is congruent with results of Near et al. (2013).
Comment: Greenwood et al. (1966) hypothesized a close affinity between Drepane and ephippids.
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.
Comment: Although support for this clade is weak (50%) in our analysis, it has been consistently obtained by previous studies with higher nodal support (90-99% in Near et al. (2012a) and 70-89% in Near et al. (2013)).
Order Lophiiformes (100%). This order is the sister group of Tetraodontiformes (55% bootstrap); also supported by anatomical evidence (Chanet et al., 2013), larval characters (Baldwin, 2013), and previous molecular studies (e.g. Dettaï and Lecointre, 2008; Miya et al., 2003; Miya et al., 2010).
Not examined: Brachionichthyidae, Lophichthyidae, Tetrabrachiidae.
Not examined: Caulophrynidae, Centrophrynidae, Diceratiidae, Linophrynidae, Neoceratiidae, Thaumatichthyidae.
Order Tetraodontiformes (100%). This order is the sister group of Lophiiformes (55% bootstrap); also supported by anatomical evidence (Chanet et al., 2013), larval characters (Baldwin, 2013), and previous molecular studies (e.g. Dettaï and Lecointre 2008; Miya et al., 2003; Miya et al., 2010).
Comment: This subordinal classification differs from that proposed by Santini and Tyler (2003).
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.
Comment: inclusion of Enoploside in this suboorder differs from results obtained by Lavoué et al. (2014).
Not examined but expected affinity (Burridge and Smolenski, 2004; Greenwood, 1995): Aplodactylidae, Chironemidae, Latridae.
Comment: percichthyoids and Percichthyidae sensu Johnson (1984) are not monophyletic: the Australian species Percalates colonorum and Percalates novemaculeata are not closely related to other members of Percichthyidae (Betancur-R. et al. (2013a); Chen et al. (2014b); Lavoué et al. (2014)), so these species are herein placed in their own suborder (Peter Unmack pers. comm.; Lavoué et al. (2014)). Percalates is listed as a junior synonym of Macquaria by Eschmeyer (2014), but the type species of Macquaria (M. australasica) is closely related to other species of Macquaria (M. ambigua) within Percichthyidae sensu stricto, hence both names are valid genera (Peter Unmack et al., pers. comm.; Lavoué et al. (2014)). Percichthyidae sensu stricto includes Percilia (formerly placed in its own family Perciliidae).
Comment: Formal description of a new family for Percalates is required to comply with the ICZN.
Comment: Although the family name Cirrithidae Macleay 1841 is older than Centrarchidae Bleeker 1859, we retain the name Centrarchiformes for this order in agreement with previous usage but expand its membership following recent proposals by Near et al. (2013) and Chen et al. (2014b) and Lavoué et al. (2014).
Not examined (10 families traditionally placed in Scorpaeniformes): Apistidae, Aploactinidae, Congiopodidae, Eschmeyeridae, Gnathanacanthidae, Neosebastidae, Pataecidae, Perryenidae, Plectrogeniidae, Zanclorhynchidae.
Comment: Lautredou et al. (2013) using seven nuclear markers obtained a clade uniting Percidae and Trachinidae with full support.
Comment: We have chosen to recognize clades within this suborder as infraorders, adopting the ending "–ales" for this rank. Gasterosteales and Zoarcales are probably sister-groups (although not in our results); they have been grouped as Zoarciformes by Li et al. (2009).
Comment: Hexagrammidae as formerly defined is not monophyletic. We now split it into two families (formerly subfamilies): Hexagrammidae (sensu stricto) and Zaniolepidoales following Washington et al. (1984), Shinohara (1994), and Smith and Busby (2014). As in previous cottoid classifications, these families are placed in their own infraorders (note that previous classifications use suborders instead of infraorders).
Not examined: Jordaniidae (not listed by Eschmeyer (2014); newly defined by Smith and Busby (2014)), Ereuniidae, Normanichthyidae, and Rhamphocottidae.
Comment: Smith and Busby (2014) changed the membership of Cottidae and Psycholutridae to achieve reciprocal monopyhyly of these families. Our phylogenetic results do not resolve a monophyletic Cottidae even under the new circumscription proposed by these authors. We refer the readers to that study for more details.
This classification (version 3), as its previous versions, also buids on Wiley and Johnson (2010) and Betancur-R. et al. (2013a), intending to preserve names and taxonomic composition of groups as much as possible. However, adjustments have been made to recognize well-supported molecular clades, many of which also have been obtained by previous molecular studies (several examples discussed below--references are cited when decisions are based on other publications). Order-level or supraordinal taxa have been erected (listed as new) or resurrected on the basis of well-supported clades only (>90% bootstrap values). Current taxon names supported by previous molecular or morphological studies have been retained if congruent with our results, even if bootstrap support is low (e.g., Osteoglossocephalai sensu Arratia (1999) with only 38% bootstrap). In some cases, ordinal or subordinal taxa that were not monophyletic in our analysis are also validated, as long as the incongruence is not supported by strong bootstrap values. Examples include the suborder Blennioidei (not monophyletic here but monophyletic in Wainwright et al., 2012) and the order Pleuronectiformes (not monophyletic here but monophyletic in Betancur-R. et al., 2013b). The classification is presented in phylogenetic order up to the subordinal rank (following the branching order in our results), but families within orders (or suborders) are listed alphabetically.
A total of 67 orders are classified, of which one is new (Chaetodontiformes) and two were sinked (Terapontiformes and Cirrhitiformes) in version 3. Three orders were new in the version 1 (Holocentriformes, Istiophoriformes, and Pempheriformes) and two were new in version 2 (Lobotiformes and Terapontiformes). The ordinal status of several percomorph families examined (as well as many others unexamined) belonging to the Series Carangimorpharia, Ovalentaria, and Percomorpharia remains uncertain (incertae sedis) due to poor phylogenetic resolution. We therefore list these families as incertae sedis within each of these groups (Carangimorpharia, Ovalentaria, and Percomorpharia) awaiting new phylogenetic evidence to clarify their ordinal status.
Family names for bony fishes are based on Eschmeyer and Fong (2014) and van der Laan et al. (2013), with minor modifications, indicted in each case in the classification. Van der Laan et al. (2013) and Eschmeyer (2014) should be consulted for authorship of family names and Wiley and Johnson (2010) for authorship of ordinal and subordinal names. Our list is not intended as a comprehensive revision of valid family names. Instead, it is simply an adaptation of their lists based on published studies that we know validate or synonymize taxa using explicit phylogenetic evidence. In order to minimize the number of non-monophyletic taxa, we have changed the membership of many families traditionally recognized in ichthyology whose validity is strongly challenged by molecular/morphological phylogenetic analyses. For instance, we no longer recognize families such as Scaridae, Caesionidae, and Microdesmidae (lumped with Labridae, Lutjanidae, and Gobiidae, respectively). Accordingly, the number of non-monophyletic families decreased from 41 in version 2 to 30 in this version (indicated in each case).
A total of 496 families of bony fishes are now recognized (excluding tetrapods), of which 394 (79.4%) were examined. For each order/suborder we list all families examined as well as the unexamined families whose taxonomic affinity is expected on the basis of traditional taxonomy or phylogenetic evidence. The list of 102 unexamined families can be easily obtained from this spreadsheet that also contains the complete classification, and is intended as a resource to help fish systematists direct future sequencing efforts.
The new classification scheme presented here (version 3) should be considered work in progress as any other hypothesis. It is likely to include involuntary errors and omissions in addition to the many unexamined, sedis mutabilis, and incertae sedis taxa. Updates should be forthcoming as new evidence becomes available and feedback from experts help refine it. Please send comments or concerns to classification@deepfin.org. For the most updated version always visit DeepFin.
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 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 (citations???).
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).
Gaidropsarinae is erected to family, so Lotidae is no longer paraphyletic (but see Roa-Varón and Ortí, 2009).
Cottoidei was recently revised by Smith and Busby (2014) who proposed a revised classification for all families within this suborder except the Aulorhynchidae, Gasterosteidae, and Hypotychidae (infraorder Gasterosteales), that were not examined. We follow these authors to make the following changes in the classification: (1) Hexagrammidae is split it into two families, defined as subfamilies by previous authors (e.g., Washington et al. 1984; Shinohara 1994): Hexagrammidae (sensu stricto) and Zaniolepididae. Both families are now placed in their own infraorders (Zaniolepidoales and Hexagrammales), following previous cottoid classifications. Note that these classifications use suborder instead of infraorders as presented here. (2) We now recognize the cottoid families Jordaniidae (not examined) and Scorpaenichthyidae.
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