SILURIFORMES

Datovo, Aléssio & Vari, Richard P., 2014, The adductor mandibulae muscle complex in lower teleostean fishes (Osteichthyes: Actinopterygii): comparative anatomy, synonymy, and phylogenetic implications, Zoological Journal of the Linnean Society 171 (3), pp. 554-622 : 589-596

publication ID	https://doi.org/ 10.1111/zoj.12142
persistent identifier	https://treatment.plazi.org/id/03DB6116-530D-A221-FEE4-D04C20F97EF5
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scientific name	SILURIFORMES
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SILURIFORMES View in CoL View at ENA

Descriptions

Nematogenys inermis ( Datovo & Bockmann, 2010: figs 1, 2)

All identified facial sections are partially continuous with each other, albeit with differentiation between them perceptible along specific regions of the muscle. The ricto-malaris and stegalis are separated from each other posterodorsally where the levator arcus palatini transits between them. The ricto-stegalis originates solely from the hyomandibula and preopercle in two examined specimens. A third examined specimen exhibits a very subtle tendinous connection between the origin of the malaris and the sphenotic and suprapreopercle. The rictalis and malaris remain continuous with each other medially, but are partially separated laterally. The rictalis inserts musculously on the lateral region of the dentary and angulo-articuloretroarticular. The lateral-most fibres of the malaris join the rictalis anteriorly to insert on the dorsolateral region of the coronoid process of both the dentary and angulo-articulo-retroarticular. The medial fibres of the malaris merge anteriorly with the stegalis before converging onto an insertional tendon that attaches to the posteromedial rim of the coronoid process of the dentary.

The more anterior origin of the stegalis allows its ready recognition from the ricto-malaris from a medial view. The entire stegalis arises from the quadrate, metapterygoid, hyomandibula, prootic, and sphenotic. Dorsally, the portion of the stegalis, which presumably corresponds to the epistegalis, originates medial to the levator arcus palatini from both the anterodorsal laminar portion of the hyomandibula and the neurocranium. The portion of the muscle lying anteroventral to the levator arcus palatini and with an origin solely on the suspensorium apparently corresponds to the substegalis. Nevertheless, no obvious differentiation between the epistegalis and substegalis is apparent. Anteriorly, a lateral set of fibres of the substegalis passes onto the outer surface of the segmentum facialis and broadly merges with the rictalis to jointly insert on the lateral surface of the dentary. Most fibres of the substegalis continue, however, along the inner surface of the segmentum facialis. This set of fibres joins with the epistegalis to insert musculously on the medial surfaces of the angulo-articuloretroarticular and coronomeckelian. The dorsal-most portions of the epistegalis merge anteriorly with the dorsomedial fibres of the malaris and converge onto the insertional tendon that is anchored to the posteromedial rim of the coronoid process of the dentary.

The ramus mandibularis trigeminus nerve passes medial to the bulk of the segmentum facialis and then lateral to the insertional tendon common to the dorsal portions of the malaris and stegalis.

The segmentum mandibularis is absent.

Silurichthys hasseltii ( Fig. 18 View Figure 18 )

The ricto-malaris has a broad origin from the quadrate, hyomandibula, preopercle, suprapreopercle, sphenotic, and pterotic. Slightly anterior to their common origin, the rictalis and malaris partially separate from each other along their dorsolateral portions. The rictalis inserts onto the dorsal region of the angulo-articuloretroarticular, whereas the malaris attaches to a flattened, thin intersegmental aponeurosis. This intersegmental aponeurosis is not obviously differentiated into mandibular and meckelian tendons, although the anterior portion of the aponeurosis serves as the site of origin for the segmentum facialis and the posterior portion attaches to the coronomeckelian and angulo-articulo-retroarticular.

The ramus mandibularis trigeminus passes between the rictalis and malaris.

The stegalis is completely subdivided into a lateral stegalis externa and a medial stegalis interna, both of which have their posterodorsal portions situated medial to the levator arcus palatini (see Remarks below concerning justifications for the identifications of these sections). The stegalis externa originates from the frontal, sphenotic, and hyomandibula and inserts primarily on the intersegmental aponeurosis. The ventral portion of the stegalis externa, however, merges anteriorly with the rictalis and inserts musculously on the anguloarticulo-retroarticular. A few ventrolateral-most fibres of the stegalis externa are visible from a lateral view.

The stegalis interna originates tendinously from the prootic and musculously from the anterior border of the hyomandibula. Anteriorly this section inserts tendinously on the posterior region of the maxilla proximate to the site of attachment of the maxillary barbel.

The segmentum mandibularis arises from the anterior portion of the intersegmental aponeurosis and inserts on the dentary, angulo-articulo-retroarticular, and Meckel’s cartilage. The segmentum mandibularis is undifferentiated into sections along its entirety.

Remarks

Members of the Siluriformes usually have a considerable degree of continuity between the rictalis and malaris. These sections may be differentiated from each other by a partial separation along their lateral portions ( Fig. 18 View Figure 18 ; Juge, 1898; Munshi, 1960; Diogo & Chardon, 2000a) and/or by different superficial patterns of fibre orientation ( Adriaens & Verraes, 1996; Cabuy et al., 1999; Devaere et al., 2001; Herrel et al., 2002; Schaefer & Provenzano, 2008). In some in- stances (e.g. Trichomycteridae ), the differentiation between the rictalis and malaris is subtle and limited only to the region proximate to their insertions ( Datovo & Bockmann, 2010). A completely undifferentiated and undivided ricto-malaris is present in other groups in the order, including some taxa in the Loricariidae , Plotosidae , Siluridae , and Trichomycteridae ( Takahasi, 1925; Bornbusch, 1995; Geerinckx et al., 2007; Geerinckx, Huysentruyt & Adriaens, 2009; Datovo & Bockmann, 2010). The rictalis is invariably inserted on the lower jaw, usually onto the posterior region of the coronoid process ( Takahasi, 1925). In the loricariid Lithogenes , the rictalis is partially subdivided into an endorictalis and an ectorictalis ( Schaefer & Provenzano, 2008).

In at least some members of most families of the Siluriformes , the origin of the malaris expands dorsally onto the neurocranium ( Fig. 18 View Figure 18 ) and in more derived conditions the dorsally enlarged contralateral malares contact each other midsagittally ( McMurrich, 1884; Juge, 1898; Takahasi, 1925; Eaton, 1948; Alexander, 1965; Howes, 1985b; Grande & Lundberg, 1988; Lundberg, Bornbusch & Mago-Leccia, 1991; de Pinna & Vari, 1995; Adriaens & Verraes, 1996; Cabuy et al., 1999; Devaere et al., 2001, 2006; Herrel et al., 2002; Rodiles-Hernández et al., 2005; Sarmento-Soares & Porto, 2006; de Pinna, Ferraris & Vari, 2007; Shibatta, Muriel-Cunha & de Pinna, 2007; Datovo & Bockmann, 2010). The malaris inserts on the buccopalatal membrane plus the maxilla in the Callichthyidae (see below) and on the lower jaw in both all other examined siluriforms and those for which the muscle was described in the literature ( Fig. 18 View Figure 18 ). In some cases where a distinction between the malaris and rictalis is not evident from a lateral view, the malaris can be identified by its anterior association with the mandibular tendon and the segmentum mandibularis ( Fig. 18 View Figure 18 ; Takahasi, 1925; Datovo & Bockmann, 2010).

The posterodorsal portion of the stegalis almost always originates medial to the levator arcus palatini ( Fig. 18 View Figure 18 ). The origin of this section extends to the neurocranium at least in some species in the Clariidae , Nematogenyidae , Plotosidae , and Siluridae (present study; Takahasi, 1925; Adriaens & Verraes, 1996; Cabuy et al., 1999; Devaere et al., 2001, 2006; Herrel et al., 2002; Datovo & Bockmann, 2010). Anteroventral to the levator arcus palatini, the stegalis is often continuous with, and sometimes almost indistinguishable from, the remaining facial sections.

Identification of the stegalis may be even more problematic because most siluriforms exhibit significant changes in the osteological landmarks typically useful for the identification of the origin and insertion of the stegalis across most teleosts. The hyomandibula of the Siluriformes bears a prominent anterodorsal laminar outgrowth (less pronounced in the Diplomystidae and Nematogenyidae ) that extends into the area typically occupied by the metapterygoid in other teleosts (authors’ pers. observ.; Fink & Fink, 1981; Arratia, 1992). In turn, the metapterygoid is anteriorly displaced along the suspensorium, consequently assuming the position typically occupied by the endopterygoid and ectopterygoid elsewhere in the Teleostei. As a consequence of these modifications, the stegalis, which usually arises primarily from the metapterygoid in most nonsiluriform teleosts, may arise largely or solely from the hyomandibula in some siluriforms (e.g. derived trichomycterids; Datovo & Bockmann, 2010). Additionally, the coronomeckelian, which typically serves as an insertion site (via the meckelian tendon) for at least part of the stegalis across the Teleostei, is absent in several siluriform taxa ( de Pinna, 1993). To further complicate matters, parts of the stegalis may become dissociated from the remainder of that section and become attached to the maxilla, which is reduced to a small, toothless ossification supporting the maxillary barbel across all the Siluriformes except for the Diplomystidae and † Hypsidoridae ( Alexander, 1965; Grande, 1987), which resolve as the basal-most members of the order under morphological analyses. This muscle division, which serves to adduct the maxillary barbel, has been designated the adductor tentaculi ( McMurrich, 1884; Takahasi, 1925; Eaton, 1948; Alexander, 1965) or, more commonly, the retractor tentaculi ( Mahajan, 1971; Howes, 1983; Schaefer, 1990, 1997; Bornbusch, 1995; Adriaens & Verraes, 1996; Cabuy et al., 1999; Devaere et al., 2001, 2006; Herrel et al., 2002; Sarmento-Soares & Porto, 2006; Huysentruyt, Geerinckx & Adriaens, 2007; Geerinckx et al., 2009; Huysentruyt, Brunain & Adriaens, 2009). Analysis reveals, however, that different parts of the stegalis have apparently independently become attached to the maxilla in different lineages during the evolution of the Siluriformes . Thus, as is discussed further below, the retractor tentaculi as now utilized is not homologous across the Siluriformes .

In the Ictaluridae and Sisoridae , the part of the segmentum facialis that inserts on the maxilla and the buccopalatal membrane originates from the dorsal region of the hyomandibula, being the only muscle portion situated medial to the levator arcus palatini ( McMurrich, 1884; Eaton, 1948; Mahajan, 1971). The remainder of the segmentum facialis in these families originates completely lateral and ventral to the levator arcus palatini. Therefore, the section attached to the maxilla in the Ictaluridae and Sisoridae apparently corresponds to the epistegalis. Association of the epistegalis with the maxilla or buccopalatal membrane also occurs in some trichomycterids (= A3″ of Datovo & Bockmann, 2010). Both the epistegalis and the malaris insert on the maxilla and buccopalatal membrane in the Callichthyidae . The name retractor tentaculi has been used in this family to refer solely to the epistegalis ( Howes, 1983; Huysentruyt et al., 2007, 2009) or to both the epistegalis and malaris ( Geerinckx et al., 2009). In the Loricariidae , Astroblepidae , and Scoloplacidae , the epistegalis is attached to the premaxilla (= retractor premaxillae of Howes, 1983; Schaefer, 1990, 1997; Geerinckx et al., 2009). The substegalis of loricariids is further divided into an inner substegalis interna and an outer substegalis externa. According to Geerinckx et al. (2009), these subsections are derived from the undivided substegalis present earlier in ontogeny. In adult loricariids, the substegalis interna (= retractor palatini of Howes, 1983; Schaefer, 1990, 1997; = retractor veli of Geerinckx et al., 2009) becomes attached anteriorly to the buccopalatal membrane, whereas the substegalis externa retains its lower jaw attachment jointly with the ricto-malaris. Munshi (1960) also reported what seems to be a substegalis divided into interna and externa subsections in Sperata aor (Bagridae) , although in this species both subdivisions retain an insertion on the lower jaw.

Determination of the identity of the so-called retractor tentaculi is more complicated amongst other groups in the Siluriformes , such as the Clariidae and Siluridae . In these families, the part of the segmentum facialis that inserts on the lower jaw is posteriorly divided into two major portions: the first portion lying lateral (= ricto-malaris) and the second medial to the levator arcus palatini ( Fig. 18 View Figure 18 ; Juge, 1898; Takahasi, 1925; Bornbusch, 1995; Adriaens & Verraes, 1996, 1997; Cabuy et al., 1999; Devaere et al., 2001, 2006; Herrel et al., 2002). This medial portion of the segmentum facialis has a broad origin that involves the hyomandibula and sometimes also the metapterygoid, quadrate and neurocranium. The retractor tentaculi of clariids and silurids lies completely medial to this inner facial division and also has a broad origin that may also involve the hyomandibula, metapterygoid, quadrate, and neurocranium. Thus, it appears that the entire stegalis in the Clariidae and Siluridae has been subdivided along a parasagittal plane, thereby forming two subsections provisionally named the stegalis externa and stegalis interna ( Fig. 18 View Figure 18 ). This division, therefore, apparently does not correspond to a separation of the stegalis into the dorsal epistegalis and ventral substegalis as found elsewhere in the Teleostei. Homology hypotheses for the retractor tentaculi of the Clariidae and Siluridae must be viewed as tentative because they are based upon a comparatively sparse sampling within the hyperdiverse Siluriformes . The identity of the so-called retractor tentaculi of other siluriform groups is even more uncertain given the limited available detailed information on this muscle system in the literature. Additional research centred on the adductor mandibulae is prerequisite for a clarification of the identities of these facial sections across the expanse of the Siluriformes .

As amply documented by Geerinckx et al. (2009), and confirmed by our observations, the muscle designat- ed as the retractor tentaculi or muscle ‘a’ in the Loricariidae , Astroblepidae , and Scoloplacidae ( Howes, 1983; Schaefer, 1990, 1997; Geerinckx et al., 2007) is actually derived from the extensor tentaculi rather than the adductor mandibulae. For this reason, Geerinckx et al. (2009) proposed renaming this muscle as the levator tentaculi. A muscle similarly named the retractor tentaculi by Diogo, Chardon & Vandewalle (2003d) and Sarmento-Soares & Porto (2006) in the auchenipterid Centromochlus heckelii similarly seems to be clearly derived from the extensor tentaculi, inasmuch as these two muscles are in close contact with each other and given that the supposed retractor tentaculi originates from the anterior portion of the neurocranium rather than from the suspensorium.

As detailed above, the term retractor tentaculi has been applied to at least five different entities: the stegalis interna ( Clariidae and possibly Siluridae ), epistegalis ( Callichthyidae , Ictaluridae , and Sisoridae ), malaris and epistegalis jointly ( Callichthyidae ), a dorsal subdivision of the extensor tentaculi ( Loricariidae ), and a ventral subdivision of the extensor tentaculi ( Auchenipteridae ). Owing to this confusing and ambiguous application of the term retractor tentaculi and given that this name in isolation fails to indicate the homologies of the muscle with the corresponding muscle section of other teleosts, we prefer to avoid the use of this term. Use of distinct names for other sections that do not attach to the lower jaw in other siluriform taxa (e.g. retractor premaxillae and retractor veli of derived taxa in the Loricarioidea) should in our opinion be similarly avoided given that the homologies of these muscles were recently clarified ( Geerinckx et al., 2009).

Many members of the Siluriformes completely lack a segmentum mandibularis. When present ( Fig. 18 View Figure 18 ), this segment is often small and has its origin limited to the intersegmental aponeurosis ( Takahasi, 1925).

Descriptions of, and nomenclature for, the adductor mandibulae sections in some studies dealing with units in the Siluriformes ( Alexander, 1965; Schaefer & Lauder, 1986; Wu & Shen, 2004) lack sufficient detail for homology determinations. Consequently, these publications are not included in the synonymy below.

In a series of at least 28 studies published between 1999 and 2007, 26 of them as sole or first author, Diogo discussed the pectoral and superficial cranial muscles – including the adductor mandibulae complex – of many groups across the breadth of the Siluriformes ( Diogo, Vandewalle & Chardon, 1999; Diogo & Chardon, 2000a, b; Diogo, Oliveira & Chardon, 2000, 2001; Diogo, Chardon & Vandewalle, 2001, 2002, 2003a, b, c, d, e, 2004a, b, c, d, 2006a; b; Oliveira et al., 2001, 2002; Diogo, 2002, 2004a, b, 2005, 2007a, b; Diogo & Vandewalle, 2003; Diogo & Bills, 2006). As a consequence, the Siluriformes ranks very high amongst orders within the Teleostei in terms of numbers of myological studies. Notwithstanding this impressive number of analyses, these papers did not yield a better understanding of the adductor mandibulae across the order. As an example, Datovo & Bockmann (2010) documented, and our analysis confirms, numerous discrepancies between the observations made in the adductor mandibulae of Nematogenys inermis – the single extant member of the Nematogenyidae – and the morphology of the muscle reported by Diogo et al. (2006a). Problematic aspects of the study by Diogo et al. (2006a) include: inaccuracies in the illustrations and morphological descriptions, misidentification of muscle sections, and an unclear and apparently inconsistent criteria for the recognition of muscle sections (see the detailed analysis in Datovo & Bockmann, 2010: 231, 232). In so far as data as to the morphology of the adductor mandibulae of Nematogenys inermis are congruent across all publications by other authors based on multiple examined specimens (present study; Howes, 1983; Datovo & Bockmann, 2010), it is unlikely that intraspecific variation accounts for the highly divergent data of Diogo et al. (2006a). Shortcomings in Diogo et al.’s (2006a) accounts of the adductor mandibulae in Nematogenys inermis , furthermore, are not an isolated case. Rather, personal observations on the morphology of the adductor mandibulae of other siluriforms treated in other publications by Diogo (and in some instances co-authors) demonstrate similar problems.

Further complicating the issue is the fact that the nomenclature for the facial sections of the adductor mandibulae originally proposed in Diogo & Chardon (2000a), and employed thenceforth in all publications by Diogo, fails to reflect the homologies of the muscle sections amongst different orders in the Teleostei. The term A2, for example, was applied in that series of papers to five different muscle sections ( Datovo & Vari, 2013: 19). More notably, the terminology proposed by Diogo & Chardon (2000a) fails to reflect the muscle homologies even across different taxa within the Siluriformes . Reflective of this is the fact that although Diogo & Chardon (2000a) recognized that his retractor tentaculi ‘is derived from the inner [facial] section of the adductor mandibulae (A3)’, this homology is implicitly ignored in most of the subsequent studies by Diogo. For example, the adductor mandibulae of the pimelodoid Pimelodus blochii is described as having five sections – A1-ost, A2, A3′-d, A3′-v, and A3″ – and no retractor tentaculi in Diogo (2005). The muscle of a second pimelodoid, Heptapterus mustelinus , is described in an almost identical repetition of the muscle description of Pimelodus blochii , with the same five subdivisions, all exhibiting the same basic characteristics; however, in Heptapterus mustelinus a retractor tentaculi is reported as present ( Diogo, 2007a). Consequently, in so far as the retractor tentaculi is considered a subdivision of the A3 ( Diogo & Chardon, 2000a), at least one of the names A3′-d, A3′-v, or A3″ is inconsistently applied to nonhomologous structures in Pimelodus blochii and Heptapterus mustelinus . Similar issues apply across nearly all of the publications on siluriforms by Diogo. In light of the scale of the problems in terms of anatomical ac- curacy, explicitness of data presentation, interpretation of observations, and logical consistency of the publications authored by Diogo, it proved impossible to include the copious amount of data generated by that author into our synonymy or utilize that information for phylogenetic analyses.

Synonymy

Segmentum facialis

A2A3: Datovo, Carvalho & Ferrer (2012): Trichomycterus .

Pars ricto-malaris

?A 1 [sic]: Howes (1983): Callichthys , Hoplosternum .

A1-OST plus A2A3′: Adriaens, Baskin & Coppens (2010): Trichomycterus .

A 2: Bornbusch (1995): Belodontichthys , Ceratoglanis , Hemisilurus , Hito (= Pterocryptis ), Kryptopterus , Kryptopterus apogon (= Phalacronotus apogon ), Kryptopterus bleekeri (= Phalacronotus bleekeri ), Kryptopterus hexapterus (= Micronema hexapterus ), Kryptopterus micronema (= Phalacronotus micronemus ), Kryptopterus moorei (= Micronema moorei ), Kryptopterus parvanalis (= Phalacronotus parvanalis ), Ompok , Ompok eugeneiatus (= Kryptopterus eugeneiatus ), Ompok sabanus (= Kryptopterus sabanus ), Pterocryptis , Silurichthys , Silurus , Wallago .

A2: Datovo & Bockmann (2010): Bullockia , Copionodon , Haemomaster , Hatcheria , Homodiaetus , Ituglanis , Listrura , Microcambeva , Nematogenys , Ochmacanthus , Paracanthopoma , Parastegophilus , Paravandellia , Pareiodon , Pseudostegophilus , Sarcoglanis , Scleronema , Stauroglanis , Trichogenes , Trichomycterus , Tridentopsis , Vandellia ; Schaefer & Provenzano (2008): Lithogenes .

A2β: Wu & Shen (2004): Parasilurus (= Silurus ).

A 2 A′ 3: Adriaens & Verraes (1996): Clarias ; Adriaens & Verraes (1997): Clarias ; Takahasi (1925): Fluvidraco [sic] nudiceps (=? Tachysurus nudiceps or? Tachysurus fulvidraco ), Liobagrus , Parasilurus (= Silurus ), Plotosus , Pseudobagrus .

A 2 A 3 ′: Cabuy et al. (1999): Clariallabes , Clarias , Gymnallabes ; Devaere et al. (2001): Channallabes ; Devaere et al. (2006): Platyclarias ; Herrel et al. (2002): Channallabes , Clariallabes , Clarias , Gymnallabes .

External part: Geerinckx et al. (2009): Ancistrus ; Geerinckx et al. (2007): Ancistrus .

Portion superficielle or Am 1: Juge (1898): Silurus .

Pars rictalis

A 1 or lateral fibres of muscle ‘b’: Howes (1983): Ancistrus , Astroblepus , Chaetostoma , Cochliodon (= Hypostomus ), Farlowella , Hemiancistrus, Hemiodonichthys [sic] (= Hemiodontichthys ), Hemipsilichthys calmoni (= Pareiorhaphis cameroni ), Hypoptopoma carinata (= Oxyropsis carinata ), Hypostomus , Hypostomus spinossisimus [sic] (= Isorineloricaria spinosissima ), Ixinandria , Lipopterichthys , Lithoxus , Loricaria , Loricaria brunneus (= Loricariichthys brunneus ), Loricaria jubata ( Rineloricaria jubata ), Loricaria labialis (= Loricariichthys labialis ), Loricaria lanceolata (= Rineloricaria lanceolata ), Loricaria microlepidogaster (= Rineloricaria microlepidogaster ), Loricaria platystoma (= Cteniloricaria platystoma ), Loricaria strigilata (= Rineloricaria strigilata ), Loricaria teffeana (= Rineloricaria teffeana ), Loricaria variegata (= Crossoloricaria variegata ), Loricaria venezeuelae ( Crossoloricaria venezuelae ), Metaloricaria , Otocinclus nigricauda (= Hisonotus nigricauda ), Panaque , Peckoltia pulcher ( Dekeyseria pulchra ), Pogonopoma , Pogonopomoides (= Pogonopoma ), Pseudancistrus , Pseudohemiodon , Pterygoplichthys, Rhadniloricaria [sic] (= Rhadinoloricaria ), Stoniella [sic] (= Pseudacanthicus ), Sturisoma .

A 1 OST: Huysentruyt et al. (2007): Corydoras ; Huysentruyt et al. (2009): Corydoras .

A1-ost: Sarmento-Soares & Porto (2006): Centromochlus , Glanidium , Tatia .

A1-OST plus A2A3′β: Adriaens et al. (2010): Trichomycterus .

A2′: Datovo & Bockmann (2010): Bullockia , Haemomaster , Hatcheria , Homodiaetus , Ituglanis , Listrura , Nematogenys , Ochmacanthus , Paracanthopoma , Parastegophilus , Paravandellia , Pareiodon , Pseudostegophilus , Sarcoglanis , Scleronema , Trichogenes , Trichomycterus , Tridentopsis , Vandellia .

A2α: Wu & Shen (2004): Pseudobagrus taiwanensis (= Tachysurus taiwanensis ).

A 2 A′ 3 β: Adriaens & Verraes (1996): Clarias .

A 2 A 3 ′β: Cabuy et al. (1999): Clariallabes , Clarias , Gymnallabes ; Devaere et al. (2001): Channallabes ; Devaere et al. (2006): Platyclarias .

A2ventral: Schaefer & Provenzano (2008): Lithogenes .

Adductor 1 or Ad 1: Munshi (1960): Mystus aor ( Sperata aor ).

Adductor mandibulae superficialis: Mahajan (1971): Sisor .

External division: Gosline (1989): Diplomystes .

External part: Geerinckx et al. (2009): Corydoras .

Partie latérale de Am 1 or ‘a’: Juge (1898): Silurus .

Pars ectorictalis

A2ventral β: Schaefer & Provenzano (2008): Lithogenes .

Pars endorictalis

A2ventral α: Schaefer & Provenzano (2008): Lithogenes .

Pars malaris

A 2: Huysentruyt et al. (2007): Corydoras ; Huysentruyt et al. (2009): Corydoras .

A 2 or posterior fibres of muscle ‘b’: Howes (1983): Ancistrus , Astroblepus , Chaetostoma , Cochliodon (= Hypostomus ), Farlowella , Hemiancistrus, Hemiodonichthys [sic] (= Hemiodontichthys ), Hemipsilichthys calmoni (= Pareiorhaphis cameroni ), Hypoptopoma carinata (= Oxyropsis carinata ), Hypostomus , Hypostomus spinossisimus [sic] (= Isorineloricaria spinosissima ), Ixinandria , Lipopterichthys , Lithoxus , Loricaria , Loricaria brunneus (= Loricariichthys brunneus ), Loricaria jubata ( Rineloricaria jubata ), Loricaria labialis (= Loricariichthys labialis ), Loricaria lanceolata (= Rineloricaria lanceolata ), Loricaria microlepidogaster (= Rineloricaria microlepidogaster ), Loricaria platystoma (= Cteniloricaria platystoma ), Loricaria strigilata (= Rineloricaria strigilata ), Loricaria teffeana (= Rineloricaria teffeana ), Loricaria variegata (= Crossoloricaria variegata ), Loricaria venezeuelae ( Crossoloricaria venezuelae ), Metaloricaria , Otocinclus nigricauda (= Hisonotus nigricauda ), Panaque , Peckoltia pulcher ( Dekeyseria pulchra ), Pogonopoma , Pogonopomoides (= Pogonopoma ), Pseudancistrus , Pseudohemiodon , Pterygoplichthys, Rhadniloricaria [sic] (= Rhadinoloricaria ), Stoniella [sic] (= Pseudacanthicus ), Sturisoma .

A2: Sarmento-Soares & Porto (2006): Centromochlus , Glanidium , Tatia .

A2″: Datovo & Bockmann (2010): Bullockia , Haemomaster , Hatcheria , Homodiaetus , Ituglanis , Listrura , Nematogenys , Ochmacanthus , Paracanthopoma , Parastegophilus , Paravandellia , Pareiodon , Pseudostegophilus , Sarcoglanis , Scleronema , Trichogenes , Trichomycterus , Tridentopsis , Vandellia .

A 2 A′ 3 α: Adriaens & Verraes (1996): Clarias .

A 2 A 3 ′α: Cabuy et al. (1999): Clariallabes , Clarias , Gymnallabes ; Devaere et al. (2001): Channallabes ; Devaere et al. (2006): Platyclarias .

A2A3′α: Adriaens et al. (2010): Trichomycterus .

A2dorsal: Schaefer & Provenzano (2008): Lithogenes .

Adductor 2 or Ad 2: Munshi (1960): Mystus aor ( Sperata aor ).

Adductor mandibulae profundus 1 and 2: Mahajan (1971): Sisor .

Dorsolateral division of retractor tentaculi: Geerinckx et al. (2009): Corydoras .

Partie médiale de Am 1 or ‘b’: Juge (1898): Silurus .

Pars stego-malaris Internal division: Gosline (1989): Diplomystes .

Pars stegalis

A3: Datovo & Bockmann (2010): Bullockia , Copionodon , Haemomaster , Hatcheria , Homodiaetus , Ituglanis , Listrura , Microcambeva , Nematogenys , Ochmacanthus , Paracanthopoma , Parastegophilus , Paravandellia , Pareiodon , Pseudostegophilus , Sarcoglanis , Scleronema , Stauroglanis , Trichogenes , Trichomycterus , Tridentopsis , Vandellia ; Wu & Shen (2004): Parasilurus (= Silurus ), Pseudobagrus taiwanensis (= Tachysurus taiwanensis ).

Pars stegalis externa

A 3: Bornbusch (1995): Belodontichthys , Ceratoglanis , Hemisilurus , Hito (= Pterocryptis ), Kryptopterus , Kryptopterus apogon (= Phalacronotus apogon ), Kryptopterus bleekeri (= Phalacronotus bleekeri ), Kryptopterus hexapterus (= Micronema hexapterus ), Kryptopterus micronema (= Phalacronotus micronemus ), Kryptopterus moorei (= Micronema moorei ), Kryptopterus parvanalis (= Phalacronotus parvanalis ), Ompok , Ompok eugeneiatus (= Kryptopterus eugeneiatus ), Ompok sabanus (= Kryptopterus sabanus ), Pterocryptis , Silurichthys , Silurus , Wallago .

A″ 3: Adriaens & Verraes (1996): Clarias ; Takahasi (1925): Parasilurus (= Silurus ).

A 3 ″: Adriaens & Verraes (1997): Clarias ; Cabuy et al. (1999): Clariallabes , Clarias , Gymnallabes ; Devaere et al. (2001): Channallabes ; Devaere et al. (2006): Platyclarias ; Herrel et al. (2002): Channallabes , Clariallabes , Clarias , Gymnallabes .

Portion profonde or Am 2: Juge (1898): Silurus .

Pars stegalis interna

Retractor tentaculi: Adriaens & Verraes (1996): Clarias ; Adriaens & Verraes (1997): Clarias ; Bornbusch (1995): Belodontichthys , Ceratoglanis , Hemisilurus , Hito (= Pterocryptis ), Kryptopterus , Kryptopterus apogon (= Phalacronotus apogon ), Kryptopterus bleekeri (= Phalacronotus bleekeri ), Kryptopterus hexapterus (= Micronema hexapterus ), Kryptopterus micronema (= Phalacronotus micronemus ), Kryptopterus moorei (= Micronema moorei ), Kryptopterus parvanalis (= Phalacronotus parvanalis ), Ompok , Ompok eugeneiatus (= Kryptopterus eugeneiatus ), Ompok sabanus (= Kryptopterus sabanus ), Pterocryptis , Silurichthys , Silurus , Wallago ; Cabuy et al. (1999): Clariallabes , Clarias , Gymnallabes ; Devaere et al. (2001): Channallabes ; Devaere et al. (2006): Platyclarias ; Takahasi (1925): Parasilurus (= Silurus ).

Rétracteur du barbillon: Juge (1898): Silurus .

Pars epistegalis

A3″: Datovo & Bockmann (2010): Bullockia , Copionodon , Hatcheria , Ituglanis , Listrura , Microcambeva , Nematogenys , Sarcoglanis , Scleronema , Stauroglanis , Trichogenes , Trichomycterus ; Sarmento-Soares & Porto (2006): Centromochlus , Glanidium , Tatia .

Adductor 4 or Ad 4: Munshi (1960): Mystus aor ( Sperata aor ).

Adductor tentaculi: Eaton (1948): Ictalurus ; Amiurus (= Ameiurus ).

Dorsomedial division of retractor tentaculi: Geerinckx et al. (2009): Corydoras .

Retractor premaxillae: Geerinckx et al. (2009): Ancistrus ; Geerinckx et al. (2007): Ancistrus ; Schaefer & Provenzano (2008): Lithogenes .

Retractor premaxillae or muscle ‘c’: Howes (1983): Ancistrus , Astroblepus , Chaetostoma , Cochliodon (= Hypostomus ), Farlowella , Hemiancistrus, Hemiodonichthys [sic] (= Hemiodontichthys ), Hemipsilichthys calmoni (= Pareiorhaphis cameroni ), Hypoptopoma carinata (= Oxyropsis carinata ), Hypostomus , Hypostomus spinossisimus [sic] (= Isorineloricaria spinosissima ), Ixinandria , Lipopterichthys , Lithoxus , Loricaria , Loricaria brunneus (= Loricariichthys brunneus ), Loricaria jubata ( Rineloricaria jubata ), Loricaria labialis (= Loricariichthys labialis ), Loricaria lanceolata (= Rineloricaria lanceolata ), Loricaria microlepidogaster (= Rineloricaria microlepidogaster ), Loricaria platystoma (= Cteniloricaria platystoma ), Loricaria strigilata (= Rineloricaria strigilata ), Loricaria teffeana (= Rineloricaria teffeana ), Loricaria variegata (= Crossoloricaria variegata ), Loricaria venezeuelae ( Crossoloricaria venezuelae ), Metaloricaria , Otocinclus nigricauda (= Hisonotus nigricauda ), Panaque , Peckoltia pulcher ( Dekeyseria pulchra ), Pogonopoma , Pogonopomoides (= Pogonopoma ), Pseudancistrus , Pseudohemiodon , Pterygoplichthys, Rhadniloricaria [sic] (= Rhadinoloricaria ), Stoniella [sic] (= Pseudacanthicus ), Sturisoma ; Schaefer (1990): Cochliodon (= Hypostomus ), Hypostomus , Isorineloricaria , Neoplecostomus ; Schaefer (1997): Otocinclus .

Retractor tentaculi: Howes (1983): Callichthys , Hoplosternum ; Huysentruyt et al. (2007): Corydoras ; Huysentruyt et al. (2009): Corydoras ; Mahajan (1971): Sisor .

Pars substegalis

A 2: Howes (1983): Callichthys , Hoplosternum .

A′ 3: Huysentruyt et al. (2007): Corydoras .

A 3 ′: Huysentruyt et al. (2009): Corydoras .

A3′: Datovo & Bockmann (2010): Bullockia , Copionodon , Hatcheria , Ituglanis , Listrura , Microcambeva , Nematogenys , Sarcoglanis , Scleronema , Stauroglanis , Trichogenes , Trichomycterus ; Sarmento-Soares & Porto (2006): Centromochlus , Glanidium , Tatia .