A new classification of the family Ariidae (Osteichthyes: Ostariophysi: Siluriformes) based on combined analyses of morphological and molecular data Author Marceniuk, Alexandre Pires Programa de Pós-Graduação em Ciências Biológicas, Universidade Federal da Paraíba, conjunto Presidente Castelo Branco III, João Pessoa, PB 58050 - 000, Brazil & Programa de Pós-Graduação em Ecologia e Conservação, Universidade Estadual da Paraíba, Campina Grande, PB 58429 - 500, Brazil & Departamento de Biologia Estrutural e Funcional, Instituto de Biociências, Universidade Estadual Paulista Júlio de Mesquita Filho (UNESP), Botucatu, SP 18618 - 970, Brazil a_marceniuk@hotmail.com Author Oliveira, Claudio Departamento de Biologia Estrutural e Funcional, Instituto de Biociências, Universidade Estadual Paulista Júlio de Mesquita Filho (UNESP), Botucatu, SP 18618 - 970, Brazil Author Ferraris Jr, Carl J. 545 NE Floral Pl., Portland, OR 97232, USA text Zoological Journal of the Linnean Society 2024 2023-08-21 200 2 426 476 http://dx.doi.org/10.1093/zoolinnean/zlad078 journal article 296812 10.1093/zoolinnean/zlad078 a3363dd0-7f56-442a-9e91-9c936714d0ee 0024-4082 11240120 5D6C7EBF-E568-4100-9364-2DD357003878 Subfamily Ariinae Bleeker, 1858 MP 100, ML 100 ( Figs 1–2 ) Type genus: Arius Valenciennes, 1840 . Diagnosis Mesethmoid medial notch narrow and deep (1, 0> 1), state 0 in Cryptarius and Ketengus , state 2 in Batrachocephalus , Bleekeriella , Brustiarius utarus , Cochlefelis , Doiichthys , Nedystoma , Nemapteryx , Netuma , Notarius grandicassis , Osteogeneiosus , Pararius mastersi , Papuarius , Potamosilurus (except Potamosilurus velutinus ), Sciades parkeri , and Sciades proops ; bones forming cephalic shield granulated (33, 0> 1) ; vomerine tooth plates rounded (56, 0> 1), state 2 in Brustiarius (except Brustiarius utarus ) and Pararius proximus , state 3 in Aspistor ; accessory tooth plates large, oval to subtriangular (60, 0> 2), state 0 in Aceroichthys , Ariopsis guatemalensis , Bleekeriella , Chinchaysuyoa , Cochlefelis , Neoarius , Paracinetodus , and Papuarius , state 1 in Brustiarius , Hexanematichthys , Nemapteryx , Pararius , and Potamarius usumacintae , state 2 in Ariopsis (except Ariopsis guatemalensis ), Arius aff. nenga , Arius dispar , Arius maculatus , Arius oetik , Carlarius , Genidens machadoi , Netuma , Notarius , Occidentarius , Pseudosciades , and Sciades , state 3 in Batrachocephalus , Cathorops , Cephalocassis , Cryptarius , Doiichthys , and Hemiarius ; distance between optic foramen and trigemino-facialis foramen about twice as large as trigemino-facialis foramen ( 68, 0> 1) , state 0 in Paragenidens and Potamosilurus ; articulation of autopalatine with lateral ethmoid moderately large ( 109, 0> 1) ; anterior cartilage of autopalatine moderately long, one-third to one-fifth as long as bone itself (113, 0> 1), state 0 in Osteogeneiosus and state 3 in Batrachocephalus and Ketengus ; premaxilla wide and moderately long, its length more than three times in width (120, 0> 1), state 0 in Aceroichthys , Brustiarius uterus , and Cochlefelis , state 2 in Arius gagora , Cathorops , Cephalocassis , Hemipimelodus , Neoarius hainesi , Notarius rugispinis , Notarius phrygiatus , Paracinetodus , and Plicofollis (except Plicofollis platystomus ) and state 3 in Doiichthys , Cinetodus , Pachyula , Paragenidens , and Potamarius (except Potamarius usumacintae ); anteroventral portion of opercle subtrapezoidal, moderately long (127, 0> 1), state 0 in Aceroichthys , Arius leptonotacanthus , Arius nenga , Arius oetik , Bleekeriella , Brustiarius utarus , Cinetodus , Hexanematichthys , Nedystoma , Pararius mastersi , and Pseudosciades and state 2 in Cathorops , Crypatrius , Hemiarius , and Pachyula ; more than one-half of posterior part of interopercle contacting ventral margin of opercle (131, 0> 1), state 0 in Arius leptonotacanthus , Batrachocephalus , Bleekeriella , Ketengus , Neoarius (except Neoarius hainesi ), Notarius (except Notarius rugispinis and Notarius phrygiatus ), Plicofollis platystomus , and Sciades (except Sciades couma and Sciades herzbergii ); metapterygoid as deep as long in perpendicular section (135, 0> 1), state 0 in Doiichthys , state 2 in Cathorops , Cinetodus , Chinchaysuyoa , Cryptarius , Hemipimelodus , Pachyula , Paragenidens , Plicofollis , and Potamarius izabalensis and state 3 in Batrachocephalus and Ketengus ; first external branchiostegal ray as broad proximally as distally (147, 0> 1); second basibranchial mushroom shaped (166, 0> 1), state 0 in Cochlefelis , Brustiarius , Doiichthys , Neoarius , Notarius (except Notarius rugispinis , Notarius planiceps , and Notarius phrygiatus ), Osteogeneiosus , Sciades couma , and Sciades herzbergii . Ambiguous optimization: Lateral horn of lateral ethmoid compressed and spatulate (11, 0> 1), state 0 in Aspistor , Cephalocassis , Cryptarius , Hemiarius , Hemipimelodus , Doiichthyina (except Bleekeriella , Cinetodus , Neoarius , Pachyula , Paracinetodus , Papuarius , and Potamosilurus velutinus ), Notarius (except Notarius armbrusteri and Notarius biffi ), Plicofollis platystomus , Plicofollis layardi , and Sciades (except Sciades couma and Sciades couma ); lateral horn of lateral ethmoid of variable length and lateroposteriorly oriented (12, 0> 1), state 0 in Aceroichthys , Brustiarius (except Brustiarius utarus ), Cochlefelis danielsi , Pararius , Plicofollis layardi , Sciades (except Sciades couma and Sciades herzbergii ) and state 2 in Doiichthys , Paragenidens , Plicofollis dussumieri , Plicofollis nella , Plicofollis polystaphylodon , and Potamarius (except Potamarius usumacintae ); posterior cranial fontanel relatively narrow and long (27, 0> 1), state 0 in Arius leptonotacanthus , Arius nenga , Brustiarius , Cathorops (except Cathorops dasycephalus ), Genidentini (except Paragenidens ), Neoarius hainesi , Netuma , Plicofollis (except Plicofollis platystomus ), state 2 in Aspistor , Hemiarius , Notarius rugispinis , Notarius phrygiatus , Papuarius , and state 3 in Cephalocassis , Doiichthys , Hemipimelodus , Nedystoma , and Nemapteryx ; extrascapular subquadrangular (37, 0> 1), state 0 in Arius , Batrachocephalus , Betancurichthys , Brustiarius (except Brustiarius utarus ), Genidens , Ketengus , Notarius (except Notarius rugispinis , Notarius planiceps , Notarius phrygiatus , and Notarius troschelii ), Pararius mastersi , Paragenidens , Plicofollis , Potamarius (except Potamarius izabalensis ), Sciades dowii and state 2 Cathorops , Cinetodus , and Paracinetodus ; temporal fossa moderate to very large (39, 0> 1), state 0 in Arius nenga , Bleekeriella , Brustiarius (excluded Brustiarius utarus ), Cathorops , Cryptarius , Genidens , Hemipimelodus , and Pararius ; articulation of autopalatine with lateral ethmoid slightly displaced to posterior portion of bone ( 111, 1> 2) , state 0 in Doiichthys , and state 1 in Brustiarius , Cochlefelis , Nemapteryx , and Neoarius ; dorsal crest of hyomandibula present (140, 0> 1) ; first epibranchial parallel to second epibranchial (179, 1> 0), state 1 in Ketengus ; nuchal plate and parieto-supraoccipital contacting one another through a convex-concave articulation (220, 0> 1), state 0 in Notarius armbrusteri , Notarius biffi , Notarius kessleri , and Notarius neogranatensis and state 2 in Aspistor , Notarius troschelii , Pararius mastersi , Sciades (except Sciades herzbergii , Sciades parkeri , and Sciades proops ). Included tribes Incertae sedis Ariinae Ariini Bleeker, 1858 Cathoropsini tribe nov. Genidentini tribe nov. Sciadeini tribe nov. Key to New World genera, Ariinae 4a. Adipose-fin base long or very short; ventral tip of subvertebral process acute or spatulate; otolith longer than wide; otolith antero-mesial process inconspicuous ….. Cathoropsini ...................................................................................................................... 5 4b. Adipose-fin base moderately long; ventral tip of subvertebral process rounded; otolith almost as long as wide; otolith anteromesial process conspicuous …. Incertae sedis , Genidentini and Sciadeini ....................................................................................... 6 5a. Accessory tooth plates small, oval; temporal fossa very reduced; anterior margin of otolith markedly irregular, concave; superficial ventral ossification of complex vertebra regularly arched ................................................................................. Cathorops 5b. Accessory tooth plates large, oval to subtriangular; temporal fossa moderate to very large; anterior margin of otolith straight or slightly irregular; otolith posterior margin rounded; superficial ventral ossification of complex vertebra keeled ................. .............................................................................................................................................................................................................. Notarius 6a. Posterior cranial fontanel absent; epiphyseal bar indistinct ................................................................................................................. 7 6b. Posterior cranial fontanel present; epiphyseal bar distinct ................................................................................................................... 8 7a. Mesethmoid median portion moderately large; distinct fenestra delimited by lateral ethmoid and frontal; lateral margin of premaxilla with slight concavity; Müllerian ramus distal third gently curved; females with conspicuous pad on pelvic fin … ( Sciadeini ) .......................................................................................................................................................................................... Ariopsis 7b. Mesethmoid median portion very wide; indistinct fenestra delimited by lateral ethmoid and frontal; lateral margin of premaxilla with a very conspicuous concavity; Müllerian ramus distal third straight; females without conspicuous pad on pelvic fin … ( Sciadeini ) ................................................................................................................................................................... Sciades 8a. Fenestra delimited by lateral ethmoid and frontal very small or indistinct; temporal fossa absent; subvertebral process indistinct or weakly developed … ( incertae sedis ) .................................................................................................................... Occidentarius 8b. Fenestra delimited by lateral ethmoid and frontal distinct and moderately wide; temporal fossa present; subvertebral process well developed ............................................................................................................................................................................................... 9 9a. Posterior cranial fontanel relatively narrow and long; premaxilla very wide and short, as long as wide (except Paragenidens grandoculis ); mesethmoid median portion very narrow (except Potamarius usumacintae ); lateral horn of lateral ethmoid long and posteriorly oriented (except Potamarius usumacintae ); optic foramen very large; distance between optic foramen and trigemino-facialis foramen small, equal to width of trigemino-facialis foramen .......................................................................... 10 9b. Posterior cranial fontanel reduced to a small opening; premaxilla wide and moderately long, its length more than three times in width; mesethmoid median portion moderately wide; lateral horn of lateral ethmoid variable length and lateroposteriorly oriented; optic foramen moderately large or very reduced; distance between optic foramen and trigemino-facialis foramen large, about twice as large as trigemino-facialis foramen ................................................................................................................... 11 10a. Snout very long, snout length 0.5–0.8 times in cephalic shield width at frontal area and 1.6–1.8 times in body width …. ( Sciadeini ) .................................................................................................................................................................................... Potamarius 10b. Snout moderately long, snout length 1.0–1.5 times in cephalic shield width at frontal area and 2.2–2.7 times in body width … ( Genidentini ) ..................................................................................................................................................................... Paragenidens 11a. Posterior process of cleithrum moderately long, about one-half vertical length of lateral face of cleithrum; gas bladder with single chamber; temporal fossa very reduced; teeth restricted to mesial two-thirds of dentary; pterotic mesial border with parieto-supraoccipital shorter than anterior border with sphenotic … ( Genidentini ) ................................................... Genidens 11b. Posterior process of cleithrum very long, equal to vertical length of lateral face of cleithrum; gas bladder with three chambers; temporal fossa moderate to very large; teeth restricted to mesial three-fourths of dentary; pterotic mesial border with parieto-supraoccipital longer than anterior border with sphenotic … ( Genidentini ) .......................................... Chinchaysuyoa . Ariinae incertae sedis ( Figs 1–3 ) Included genera Occidentarius Betancur-R. & Acero P., 2007. Remarks The monotypic genus Occidentarius listed here falls within the Ariinae in MP and ML analyses, but its position within the subfamily differs markedly among the different analyses. As such, it is treated here as a member of the Ariinae , but not included within either of the named tribes.