Champsodon snyderi, from Matsubara et al. (1964).

Belongs within: Neoteleostei.
Contains: Lampridiformes, Paracanthomorphacea, Polymixiiformes, Beryciformes, Holocentridae, Sparisomidae, Ophidiiformes, Batrachoididae, Mullidae, Scombrimorpharia, Gobiomorpharia, Carangimorpharia, Percomorpharia.

The Acanthomorphata are a clade of fishes in which hollow, unsegmented spines are present in the anterior part of the dorsal and anal fins.

Unstitching trachinoids
Published 19 June 2007

Historical classifications of the acanthomorph fishes have recognised the Trachinoidei as a suborder including what are known as the weevers and allies—mostly elongate, often dorsoventrally flattened, regularly benthic fish. In keeping with the benthic habitat (most species bury themselves in the sand for camouflage), the eyes are generally placed on top of the skull with the mouths pointing upwards. This suborder has been treated as including the stargazers and sand lances.

Greater weevers Trachinus draco, copyright Tomasz Sienicki.

To be honest, I’m having a hard time finding what, beyond a vague overall similarity, were the actual uniting characters for this group. Bond (1996) emphasises the jugular pelvic fins (placed under the throat), but this feature is not unique to trachinoids—the Blennioidei (blennies), Gobiesocidae (clingfishes) and Callionymoidei (dragonets) all also have jugular pelvics. Pietsch & Zabetian (1990) gave two characters—presence of a pelvic spur and pectoral radials being small, short or wide—uniting a collection of families that they referred to as the core of, but not necessarily delimiting, the Trachinoidei. Not surprisingly, all the molecular studies that have addressed the issue appear to recover the trachinoids as polyphyletic (Stankovic et al., 2005), and authors have disagreed significantly about which families should be included in the suborder. Being a trachinoid, it seems, is not so much a reality as a state of mind.

A number of trachinoids have poisonous spines, and the name “weever” for members of the Trachinidae is supposedly derived from the Anglo-Saxon word for “viper”. Uranoscopus (stargazers) also have electric organs behind the eyes—Bond (1996) credits them with producing 50 volts.

Black swallower Chiasmodon niger, from Goode & Bean (1896).

Cheimarrichthys fosteri (the torrentfish) is a freshwater fish unique to fast-running streams in New Zealand that is sometimes placed in its own family. My definite favourite among the families assigned to this suborder, however, has to be the Chiasmodontidae (black swallowers), another addition to the mesopelagic freakshow. Black swallowers get their name from their ability to distend their stomach to three times their own size and so swallow fish larger than themselves whole (memories of snork-eater-eaters…). Doubtless this is a very handy adaptation in the fairly sparse environment of the mesopelagic, where meals may be few and far between.

The bush at the top of the tree
Published 4 December 2008
Sailfish (Istiophorus albicans) feeding on sardines. Photo by Douglas Seifert.

In an earlier post, I introduced the concept of a wastebasket taxon—a dumping-place for usually largely unspecialised taxa that do not possess the distinctive characters to place them elsewhere. Sadly, the history of taxonomy has no shortage of examples of wastebaskets. Some of them have been progressively cleaned out in recent years; others remain sitting there in a jumbled heap, just daring reasearchers to have the courage (or foolishness) to take them on. Of all the enormous, ugly junk-piles in classification, however, perhaps none is more enormous or more ugly than the Perciformes.

The Perciformes, as pointed out by many a textbook, is the largest by a considerable margin of all orders of fishes, and of vertebrates for that matter. There are over 7000 species of Perciformes of all different varieties, from tiny gobies to gigantic marlin and tuna. Go down to a beach, drop a fishing line into the water, and you can almost guarantee that the first thing you pull up will be a perciform. What makes all this diversity even more of an achievement, though, is that the order Perciformes doesn’t really exist. Not in any way that actually makes sense. Not a single distinctive character unites all of the various taxa included in the Perciformes. As Gill & Mooi (2002) put it, “not only are there no synapomorphies to unite the order, it is not even diagnosable on the basis of shared primitive characters“. As the saying goes, it’s not even wrong.

The oarfish or king of the herrings, Regalecus glesne, a basal acanthomorph. Oarfishes can be over ten metres in length. Photo by Gerry Smith.

So if there’s nothing to unite the order, what is it doing there in the first place? “Perciform” fishes belong to a clade called Acanthomorpha, the monophyly of which is well-supported. Among other things, one of the most easily recognisable features of the Acanthomorpha is that the first few rays of the fins have been replaced by hard spines. Within the Acanthomorpha, a number of distinct orders are recognised. The dories of the order Zeiformes have a distinctive telescoping mouth that allows them to function as expert ambush predators. The pufferfishes and triggerfishes of the Tetraodontiformes, in contrast, have the bones of the mouth fused into a powerful beak for crushing hard prey. The flatfish of the order Pleuronectiformes have moved both eyes onto one side of their heads, and spend their days lying on their sides. But once these distinctive taxa have all been isolated—once you’ve taken out the fish that look like dories and the fish that look like puffers—you’re still left with a large pile of fish that, well, just kind of look like fish. This pile is what you label “Perciformes” and pretend that you’ve somehow done something about it. Needless to say, being the association of acanthomorphs that aren’t distinctive enough to be separated off, the Perciformes has varied on content quite considerably over the years.

The very undefined nature of the Perciformes is also exactly what makes it such a difficult phylogenetic problem to solve. The phylogeny of the Perciformes is nothing less than the phylogeny of the Acanthomorpha as a whole—to solve one, you must solve the other. With nearly 15,000 living species of acanthomorphs, that is a colossal task. Still, a number of poor, foolhardy souls have made the attempt over the years. The advent of molecular analyses has improved matters, though there’s still a lot of disagreement.

The figure above shows a Bayesian tree from a relatively recent effort (Dettai & Lecointre 2005). Support for a number of the clades found tended to be on the low side, but this paper and an earlier one (Chen et al. 2003) took the approach of analysing a number of separate datasets from different genes as well as the combined analysis, and seeing which clades were supported in which analyses (the theory being that a clade resulting from a number of different data sources is probably worth consideration even if the numerical support is not high every time). The clades with letters beside them are ones that appeared in a number of analyses.

Though not strongly supported, this analysis agrees with others (both molecular and morphological) in recovering a basal grade within the the Acanthomorpha that includes Polymixia (beardfishes), Zeiformes (dories), Gadiformes (cods) and Beryciformes (squirrelfishes). Members of the “Perciformes” fall in a clade that runs in the tree above from Pomatoschistus (Gobiidae) to Cyclopterus, but this clade also (surprise, surprise) includes members of a number of other orders, and the Perciformes are polyphyletic. Some of the recovered relationships have been suggested before. Clade N, for instance, includes the Tetraodontiformes, Caproidae (boarfishes) and Acanthuroidei (surgeonfishes and allies), and relationships have been suggested on morphological grounds between Tetraodontiformes and either one of the other two groups (though, ironically enough, never both at the same time). Some of the relationships suggested by molecular analyses have been entirely novel—clade N also includes the Lophiiformes (anglerfishes), which might have been just about the last group one would expect to be related to puffers.

The tree is also not without evidence of how far there is yet to go. Both Tetraodontiformes and Pleuronectiformes appear as polyphyletic, a result that is almost certainly complete twaddle in light of the distinctive morphological characters uniting members of these orders. The appearance of the opah Lampris in the middle of the Serranidae seems also pretty suspect to me (offhand, Lampris appears on the tree twice—I suspect the “Lampris” near the bottom of the tree is a misprint for Regalecus, the oarfish or king of the herrings). G. Nelson commented of fish phylogeny in 1989 that “recent work has resolved the bush at the bottom, but the bush at the top persists“. That bush still needs a lot of topiary.

Perciformes go bye-bye?
Published 3 January 2009
Anableps anableps, the four-eyed fish, a member of the newly-named clade Stiassnyiformes. The upper and lower halves of the eyes have different focal distances, allowing the fish to see clearly both above and below the water. Photo from here

In the section above, I referred to the unholy taxonomic mess that besets the Acanthomorpha (spiny-finned fishes), the clade that includes (among others) the majority of familiar marine fish, with a large number of acanthomorphs previously dumped in a cluster called “Perciformes”. “Perciformes”, I’ll remind you, was ichthyological code for “morphologically somewhat boring fish that aren’t distinctive enough to be called anything else”.

Just yesterday, I received notification of a new paper (Li et al. 2009) that identifies a number of the same clades recovered in papers such as Dettai & Lecointre (2005) (which is not entirely surprising because both Dettai and Lecointre are also authors on this one). Where Li et al. (2009) differs from previous studies, however, is that it actually introduces a whole bunch of new names for the clades it finds. (Sadly, they also do something that is a bit of a pet peeve of mine, what I call “stealth taxonomy”—introducing new taxa in a paper without indicating that they’re doing so in the title or mentioning their names in the abstract).

The above diagrams from Li et al. (2009) represent a supertree for the Acanthomorpha taken from the results of Li et al.‘s analyses. It’s been broken into four parts because that’s a lot of fish. You may need to click on the images to get a higher resolution picture to follow the next bit.

The clade labelled P at the bottom of the tree doesn’t get a name – some studies have found it as monophyletic, some haven’t. What does get a name is the clade A uniting the Zeiformes (dories) and Gadiformes (cods)—Zeioigadiformes. That, my friends, is not a name to attempt whilst drunk. The Beryciformes (squirrelfishes and allies) are paraphyletic in the supertree (they’re the first three clades leading towards the big P’ clade), but some studies (including one of the analyses of Li et al. themselves) still find it to be monophyletic—another watch this space moment. The Ophidiiformes (cusk-eels, etc., including the Carapidae that are famed for living inside the recta of sea cucumbers) are sister to the P’ clade, which is what recent studies have come to call Percomorpha. Batrachoidiformes (toadfishes) are sister to the remaining percomorphs, but as you can see there’s still a lot of messiness otherwise. Still, there are a few big clades that can be recognised, most notably the Q, L and X clades.

Sphyraena barracuda, now a member of the Carangimorpha. Photo from here.

The Q clade includes the Atherinomorpha, Mugilidae (mullets), Blennioidei (blennies) and, in most studies, the Cichlidae (cichlids), though in the supertree the cichlids are apparently feeling a bit stand-offish. Li et al. dub this clade the Stiassnyiformes, for reasons sadly unexplained. The L clade, which Li et al. dub the Carangimorpha, includes the big pelagic fish such as dolphins, trevally, marlins, tuna and mackerel, as well as the flatfish. Within the L clade, things are still rather messy, but this is mainly the fault of the flatfish still trying to make themselves look polyphyletic despite all morphological reason.

Unidentified icefish (Notothenioidei). Photo by Uwe Kils.

The X clade is labelled by Li et al. the Serraniformes. I’m guessing that they decided, probably wisely, that “Perciformes” carries just a little to much baggage. Notable members of this clade include the Percidae, Notothenioidei (Antarctic icefishes), Gasterosteidae (sticklebacks) and what were the Scorpaeniformes (now known to be polyphyletic).

Indostomus paradoxus, the armoured stickleback or paradox fish of south-east Asia. Photo from hwchoy.

A couple of smaller clades that rate a mention are the F and G clades. The F clade (now the Anabantiformes) includes the Anabantoidei (gouramis), Channidae (snakeheads), Indostomus (the paradox fish) and Synbranchiformes (swamp eels). The association of these taxa forms an intriguing clade of Old World tropical freshwater fish, with their diversity centred around southern Asia. The G clade includes an assortment of “trachinoid” fishes such as stargazers and torrent fish, but not the Trachinidae themselves. Fittingly, Li et al. label this group the Paratrachinoidei.

Part of the problem with dropping the idea of “Perciformes”, to date, has been that to do so would leave a massive pile of unplaced families just flopping all over the place. Hopefully, with the pile of new clade names presented to us by Li et al., a big step has been taken towards allowing us to never have to say “Perciformes” again.

Systematics of Acanthomorphata
Acanthomorphata (see below for synonymy)B-RB13
    |  i. s.: Acanthomorphorum forcallensisAS09
    |         AsineopidaeP93
    |           |--Nardoichthys francisci Sorbini & Bannikov 1991P93
    |           `--Asineops squamifrons Cope 1870P93
    |         Pharmacichthys [Pharmacichthyidae]P93
    |         Macconichthys [Mcconichthyidae]P93
    |           `--M. longipinnis Grande 1988P93
    |--Ctenothrissa [Ctenothrissidae]NE12
    |--Aulolepis [Aulolepidae]NE12
    |    `--A. typusW00
       |  `--ParacanthomorphaceaB-RB13
          `--Euacanthomorphacea [Acanthopterygii, Berycomorphi, Euacanthopterygii, Pediculati]B-RB13
               |  i. s.: AcanthopterygiorumP93
               |           |--A. circularis (Stinton 1968)P93
               |           `--A. dorsetensis (Stinton 1968)P93
                  |    |  i. s.: Stichoberyx polydesmus (Arambourg 1954)P93
                  |    |--HolocentridaeB-RB13
                  |    |--CaproberycidaeP93
                  |    |--AlloberycidaeP93
                  |    `--Stichocentrus [Stichocentridae]P93
                  `--Percomorphaceae (see below for synonymy)B-RB13
                       |  i. s.: Coreoperca kawamebari (Temminck & Schlegel 1843)I92
                       |         Achoerodus gouldiiG75
                       |         ExilliaFJ05
                       |         Bathyclupea [Bathyclupeidae]P01
                       |           |--B. gracilis Fowler 1938P01
                       |           |--B. malayana Weber 1913P01
                       |           `--B. megaceps Fowler 1938P01
                       |         InermiidaeB96
                       |         TrichonotidaeM58
                       |           |--TaeniolabrusR13
                       |           `--TrichonotusM58
                       |                |--T. elegans Shimada & Yoshino 1984MM09
                       |                `--T. setiger Bloch & Schneider 1801M58
                       |         SparisomidaeM58
                       |         Solenichthys [Solenichthyidae]M58
                       |           |--S. armatus (Weber 1913) [=Solenostomus armatus]M58
                       |           `--S. cyanopterus (Bleeker 1854) (see below for synonymy)M58
                       |         ScorpididaeR13
                       |           |--Medialuna californiensisR13, B96
                       |           |--Neatypus obliquus Waite 1905H90
                       |           |--AtyposomaR13
                       |           `--ScorpisS44
                       |                |--S. aequipinnis Richardso 1848C01
                       |                |--S. lineolata Kner 1865 [=S. lineolatus]C01
                       |                `--S. violaceusHS01
                       |         AsianthidaeSP03
                       |           |--Asianthus Sytchevskaya & Prokofiev 2003SP03
                       |           |    `--*A. celebratus (Daniltshenko 1968) [=Serranus celebratus]SP03
                       |           |--Pauranthus Sytchevskaya & Prokofiev 2003SP03
                       |           |    `--*P. argutulus Sytchevskaya & Prokofiev 2003SP03
                       |           `--Eosasia Sytchevskaya & Prokofiev 2003SP03
                       |                |--*E. rekubratskii Sytchevskaya & Prokofiev 2003SP03
                       |                |--E. kessleri Sytchevskaya & Prokofiev 2003SP03
                       |                `--E. lebedevi Sytchevskaya & Prokofiev 2003SP03
                       |         DichistiidaeB-RB13
                       |         Hapalogenys Richardson 1844P93 [HapalogenyidaeB-RB13]
                       |           |--H. analis Richardson 1845 [incl. H. mucronatus (Eydoux & Souleyet 1850)]M01
                       |           |--H. kishinouyei Smith & Pope 1906M01
                       |           `--H. meyeniJR10
                       |         ParascorpididaeB-RB13
                       |         Symphysanodon Bleeker 1878P93 [SymphysanodontidaeB-RB13]
                       |           |--S. maunaloaeI00
                       |           `--S. typus Bleeker 1878M58
                       |         Epigonidarum weinbergiAS09
                       |         ProtorhamphosusP93
                       |         PerciformorumP93
                       |           |--P. cepoloides Nolf & Dockery 1990P93
                       |           `--P. transitus (Sieber & Weinfurter 1967)P93
                       |         Hemilutjanus Bleeker 1876P93
                       |         GiganthiidaeP93
                       |         Caesioscorpis Whitley 1945 [Caesioscorpididae]P93
                       |           `--C. theagenes Whitley 1945H90
                       |         Trachinoideorum schwarzhansi Nolf 1988P93
                       |         Caucasichthys Bannikov, Carnevale & Parin 2011 [Caucasichthyidae]BCP11
                       |           `--*C. kuamensis Bannikov, Carnevale & Parin 2011BCP11
                       |         Erythrichthys [Erythrichthyidae]R13
                       |         Hemerocoetes [Hemerocoetidae]R13
                       |         Ditremidae [Ditremiformes]R13
                       |         Blotichthys Sorbini 1979B80
                       |           `--B. coleanus (Agassiz 1833–1844) (see below for synonymy)B80
                       |         Carangodes Heckel 1856 [incl. Arambourgella Blot 1980; Arambourgellidae, Carangodidae]B80
                       |           `--C. cephalus Heckel 1856 [=*Arambourgella cephala]B80
                             |  `--ScombrimorphariaND13
                                      `--Champsodontidae [Champsodontiformes]R13
                                           |--Centropercis nudivittis Ogilb. 1895R13
                                                |--C. atridorsalisSC07
                                                |--C. capensis Regan 1908 [incl. C. arafurensis Regan 1908]M58
                                                |--C. guentheri Regan 1908M58
                                                |--C. snyderi Franz 1910LD09
                                                `--C. vorax Günther 1867JR10

Acanthomorphata [Acanthomorpha, Anacanthini, Euacanthomorpha, Holacanthopterygii, Paracanthopterygii, Thoracici, Zeomorphi]B-RB13

Blotichthys coleanus (Agassiz 1833–1844) [=Pygaeus coleanus; incl. P. egertoni Agassiz 1833–1844, P. gibbus Agassiz 1833–1844]B80

Percomorphaceae [Aulostomoidei, Benthomorpha, Gasterosteiformes, Gasterosteoidei, Jugulares, Moronoidei, Mulloidei, Perciformes, Percoidea, Percomorpha, Percomorphi, Scleroparei, Scorpaeniformes, Thoracostei, Trachiniformes, Trachinoidei]B-RB13

Solenichthys cyanopterus (Bleeker 1854) [=Solenostoma cyanopterus; incl. Solenostoma paradoxus Kaup 1856]M58

*Type species of generic name indicated


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