Ariidae Bleeker

[[Family Ariidae Bleeker]] Identification key to genera This key is based on the use of external morphological characters allowing for the identification of taxa without the need of osteological preparations. Being artificial, it has to be used from its beginning since generic identification can be reached through different entries. 1. Maxillary barbel present............................................................................................................................ 2 - Maxillary barbel absent.................................................................................................. Batrachocephalus 2. Mental barbels present; maxillary barbel soft, fleshy along its entire length............................................ 3 - Mental barbels absent; maxillary barbel hard, bony along its entire length....................... Osteogeneiosus 3. Two pairs of mental barbels; maxillary barbel cylindrical........................................................................ 4 - A single pair of mental barbels; maxillary barbel compressed, tape-like........................................... Bagre 4. Cephalic shield clearly exposed, ornamented with grooves, ridges and granules, covered by a very thin layer of skin; anterior and median nuchal plates indistinct; posterior cleithral process and second dorsal process of cleithrum distinct and pointed.................................................................................................. 5 - Cephalic shield only slightly exposed, covered by musculature and thick skin; anterior and median nuchal plates conspicuous; posterior cleithral process and second dorsal process of cleithrum connected by a bony blade................................................................................................................................. Galeichthys 5. Premaxillary and dentary with several rows of conical, viliform or molar-like teeth............................... 6 - Premaxillary and dentary teeth with a single row of incisiform teeth.......................................... Ketengus 6. Adipose fin long, its base as long as anal-fin base.................................................................................... 7 - Adipose fin moderately long or short, its base no longer than one-half the length of anal-fin base................................. 13 7. Anterior and median nuchal plates very reduced, forming a structure of semi-lunar aspect; posterior margin of occipital process concave; occipital process triangular shaped, moderately long and wide; accessory tooth plates bearing small conical teeth..................................................................................................... 8 - Anterior and median nuchal plates very large, square to pentagonal shaped; posterior margin of occipital process convex; occipital process round, very short and wide at base; accessory tooth plates large, bearing molar-like teeth............................................................................................................................... Aspistor 8. Tooth plates associated with vomer present.............................................................................................. 9 - Tooth plates associated with vomer absent.............................................................................................. 10 9. Posterior cleithral process very short; accessory tooth plates transversely elongated and narrow.................................................................................................................................................................... Hemiarius - Posterior cleithral process moderately long; accessory tooth plates large, triangular, oval or round................................................................................................................................................................... Notarius 10. Posterior cleithral process very long; posterior cranial fontanel long and narrow.................................. 11 - Posterior cleithral process short or moderately long; posterior cranial fontanel wide and oval.............. 12 11. Accessory tooth plates present..................................................................................................... Cinetodus - Accessory tooth plates absent....................................................................................................... Pachyula 12. Fenestra limited by supraoccipital, pterotic and sphenotic present; posterior cranial fontanel very large; posterior cleithral process very short.................................................................................... Cephalocassis - Fenestra limited by supraoccipital, pterotic and sphenotic absent; posterior cranial fontanel large; posterior cleithral process moderately long....................................................................................... Amphiarius 13. Accessory tooth plates bearing large molar-like teeth............................................................................. 14 - Teeth of the accessory tooth plates small or absent, when present conical or viliform........................... 16 14. Adipose fin very short, its base less than one-half length of anal-fin base; medial groove of neurocranium limited by frontal bones and also on supraoccipital; posterior cranial fontanel very reduced or absent..................................................................................... 15 - Adipose fin moderately long, its base about one-half as long as anal-fin base; medial groove of neurocranium mostly or exclusively on supraoccipital; posterior cranial fontanel conspicuous, long and narrow................................................................................................................................................................. Arius 15. Lateral line bifurcated at caudal region; posterior cleithral process moderately long................. Plicofollis - Lateral line not bifurcated at caudal region; posterior cleithral process very short..................... Cathorops 16. Lateral line not bifurcated at caudal region............................................................................................. 17 - Lateral line bifurcated at caudal region................................................................................................... 28 17. Temporal fossa present; fontanel limited by lateral ethmoid and frontal well developed, moderate to large.................................................................................................................................................................. 18 - Temporal fossa absent; fontanel limited by lateral ethmoid and frontal very reduced or inconspicuous............................................................................................................................................................... Sciades 18. Medial groove of neurocranium limited by frontal and/or supraoccipital, rudimentary or absent............................................................................................. 19 - Medial groove of neurocranium limited by frontal and/or supraoccipital, very distinct...................................................... 22 19. Posterior cleithral process very short; posterior cranial fontanel very large........................................... 20 - Posterior cleithral process moderately long; posterior cranial fontanel moderate to large..................... 21 20. Accessory tooth plates absent..................................................................................................... Nedystoma - Accessory tooth plates present.................................................................................................... Doiichthys 21. Posterior cranial fontanel oval and well developed; mesethmoid moderately wide at region of anterior nostrils................................................................................................................................... Potamosilurus - Posterior cranial fontanel relatively long and narrow; mesethmoid very wide at region of anterior nostrils.................................................................................................................................................... Cochlefelis 22. Epioccipital not invading dorsal portion of cephalic shield.................................................................... 23 - Epioccipital invading dorsal portion of cephalic shield................................................................ Carlarius 23. Adipose fin very short, its base less than one-half length of anal-fin base.............................................. 24 - Adipose fin moderately long, its length about half as long as anal-fin base........................................... 25 24. Tooth plates associated with vomer present............................................................................... Brustiarius - Tooth plates associated with vomer absent................................................................................. Cryptarius 25. Posterior cleithral process moderately long; medial groove of neurocranium limited by frontal bones, but also on supraoccipital; posterior cranial fontanel reduced or absent or long and narrow................................................................................... 26 - Posterior cleithral process very short; medial groove of neurocranium limited mainly by supraoccipital; posterior cranial fontanel very large and oval.......................................................................... Nemapteryx 26. Posterior cranial fontanel well developed, long and narrow................................................................... 27 - Posterior cranial fontanel very reduced or absent......................................................................... Genidens 27. Tooth plates associated with vomer absent; accessory tooth plates very reduced or absent..... Potamarius - Tooth plates associated with vomer present; accessory tooth plates large.................................... Neoarius 28. Tooth plates associated with vomer absent; adipose fin moderately long, its length about half as long as anal-fin base......................................................................................................................................... Arius - Tooth plates associated with vomer present; adipose fin very short, its base less than one-half length of anal-fin base..................................................................................................................................... Netuma New classification of the Ariidae Family Ariidae Bleeker, 1862 Amissidens Kailola, 2004 Amissidens hainesi (Kailola, 2000) Amphiarius new genus Amphiarius phrygiatus (Valenciennes, 1840) Amphiarius rugispinis (Valenciennes, 1840) Arius Valenciennes, 1840 Arius acutirostris Day, 1877 - sedis mutabilis Arius africanus Günther, 1867 - sedis mutabilis Arius arenarius (Müller & Troschel, 1849) - sedis mutabilis Arius arius (Hamilton, 1822) Arius brunellii Zollezi, 1939 - sedis mutabilis Arius burmanicus Day, 1870 - sedis mutabilis Arius caelatus Valenciennes, 1840 Arius dispar Herre, 1926 Arius festinus Ng & Sparks, 2003 - sedis mutabilis Arius gagora (Hamilton, 1822) Arius jatius (Hamilton, 1822) - sedis mutabilis Arius jella Day, 1877 - sedis mutabilis Arius leptonotacanthus Bleeker, 1849 - sedis mutabilis Arius macronotacanthus Bleeker, 1846 - sedis mutabilis Arius maculatus (Thunberg, 1792) Arius madagascariensis Vaillant, 1894 Arius malabaricus Day, 1877 - sedis mutabilis Arius manillensis Valenciennes, 1840 Arius microcephalus Bleeker, 1855 - sedis mutabilis Arius nenga (Hamilton, 1822) - sedis mutabilis Arius oetik Bleeker, 1846 - sedis mutabilis Arius subrostratus Valenciennes, 1840 - sedis mutabilis Arius uncinatus Ng & Sparks, 2003 - sedis mutabilis Arius venosus Valenciennes, 1840 - sedis mutabilis Aspistor Jordan & Evermann, 1898 Aspistor luniscutis (Valenciennes, 1840) Aspistor parkeri (Traill, 1832) Bagre Cloquet, 1816 Bagre bagre (Linnaeus, 1766) Bagre marinus (Mitchill, 1815) Bagre panamensis (Gill, 1863) Bagre pinnimaculatus (Steindachner, 1876) Batrachocephalus Bleeker, 1846 Batrachocephalus mino (Hamilton, 1822) Brustiarius Herre, 1935 Brustiarius nox (Herre, 1935) Brustiarius proximus (Ogilby, 1898) - sedis mutabilis Brustiarius solidus (Herre, 1935) Carlarius new genus Carlarius gigas (Boulenger, 1911) - sedis mutabilis Carlarius heudelotii (Valenciennes, 1840) Carlarius latiscutatus (Günther, 1864) Carlarius parkii (Günther, 1864) Cathorops Jordan & Gilbert, 1822 Cathorops agassizi (Eigenmann & Eigenmann, 1888) Cathorops aguadulce (Meek, 1904) Cathorops arenatus (Valenciennes, 1840) Cathorops dasycephalus (Günther, 1864) Cathorops fuerthii (Steindachner, 1877) Cathorops hypophthalmus (Steindachner, 1877) Cathorops mapale Betancur-R. & Acero, 2005 Cathorops melanopus (Günther, 1864) Cathorops multiradiatus (Günther, 1864) Cathorops spixii (Agassiz, 1829) Cathorops steindachneri (Gilbert & Starks, 1904) Cathorops tuyra (Meek & Hildebrand, 1923) Cephalocassis Bleeker, 1858 Cephalocassis bleekeri (Popta, 1900) - sedis mutabilis Cephalocassis borneensis (Bleeker, 1851) Cephalocassis manillensis (Valenciennes, 1840) - sedis mutabilis Cephalocassis melanochir (Bleeker, 1852) Cinetodus Ogilby, 1898 Cinetodus carinatus (Weber, 1913) - sedis mutabilis Cinetodus froggatti (Ramsay & Ogilby, 1886) Cochlefelis Whitley, 1941 Cochlefelis danielsi (Regan, 1908) Cochlefelis dioctes (Kailola, 2000) - sedis mutabilis Cochlefelis insidiator (Kailola, 2000) - sedis mutabilis Cochlefelis spatula (Ramsay & Ogilby, 1886) Cryptarius Kailola, 2004 Cryptarius daugueti (Chevey, 1932) - sedis mutabilis Cryptarius truncatus (Valenciennes, 1840) Doiichthys Weber, 1913 Doiichthys novaeguineae Weber, 1913 Galeichthys Valenciennes, 1840 Galeichthys ater Castelnau, 1861 Galeichthys feliceps Valenciennes, 1840 Galeichthys peruvianus Lütken, 1874 - sedis mutabilis Genidens Castelnau, 1855 Genidens barbus (Lacépède, 1803) Genidens genidens (Cuvier, 1829) Genidens machadoi (Miranda-Ribeiro, 1918) Genidens planifrons (Higuchi, Reis & Araújo, 1982) Hemiarius Bleeker, 1847 Hemiarius hardenbergi (Kailola, 2000) - sedis mutabilis Hemiarius harmandi Sauvage, 1880 - sedis mutabilis Hemiarius stormii (Bleeker, 1858) Hemiarius sumatranus (Anonymous, 1830) Hemiarius verrucosus (Ng, 2003) - sedis mutabilis Ketengus Bleeker, 1847 Ketengus typus Bleeker, 1847 Nedystoma Ogilby, 1898 Nedystoma dayi (Ramsay & Ogilby, 1886) Nemapteryx Ogilby, 1908 Nemapteryx armiger (De Vis, 1884) Neoarius Castelnau, 1878 Neoarius augustus (Roberts, 1978) - sedis mutabilis Neoarius berneyi (Whitley, 1941) - sedis mutabilis Neoarius graeffei (Kner & Steindachner, 1867) Neoarius midgleyi (Kailola & Pierce, 1988) Neoarius pectoralis (Kailola, 2000) - sedis mutabilis Netuma Bleeker, 1858 Netuma bilineatus (Valenciennes, 1840) Netuma thalassinus (Rüppell, 1837) Notarius Gill, 1863 Notarius armbrusteri Betancur-R. & Acero, 2006 - sedis mutabilis Notarius biffi Betancur-R. & Acero, 2004 - sedis mutabilis Notarius cookei (Acero & Betancur-R., 2002) - sedis mutabilis Notarius grandicassis (Valenciennes, 1840) Notarius insculptus (Jordan & Gilbert, 1883) - sedis mutabilis Notarius kessleri (Steindachner, 1877) - sedis mutabilis Notarius lentiginosus (Eigenmann & Eigenmann, 1888) Notarius neogranatensis (Acero & Betancur-R., 2002) - sedis mutabilis Notarius osculus (Jordan & Gilbert, 1883) - sedis mutabilis Notarius planiceps (Steindachner, 1877) Notarius troschelii (Gill, 1863) Osteogeneiosus Bleeker, 1846 Osteogeneiosus militaris (Linnaeus, 1758) Pachyula Ogilby, 1898 Pachyula conorhynchus (Weber, 1913) - sedis mutabilis Pachyula crassilabris (Ramsay & Ogilby, 1886) Plicofollis Kailola, 2004 Plicofollis argyropleuron (Kuhl & van Hasselt, 1840) Plicofollis crossocheilos (Bleeker, 1846) - sedis mutabilis Plicofollis dussumieri (Valenciennes, 1840) Plicofollis magatensis (Herre, 1926) - sedis mutabilis Plicofollis nella (Valenciennes, 1840) Plicofollis platystomus (Day, 1877) - sedis mutabilis Plicofollis polystaphylodon (Bleeker, 1846) Plicofollis tenuispinis (Day, 1877) Potamarius Hubbs & Miller, 1960 Potamarius grandoculis (Steindachner, 1877) Potamarius izabalensis Hubbs & Miller, 1960 Potamarius nelsoni (Evermann & Goldsborough, 1902) Potamosilurus new genus Potamosilurus coatesi (Kailola, 1990) - sedis mutabilis Potamosilurus latirostris (Macleay, 1883) Potamosilurus macrorhynchus (Weber, 1913) Potamosilurus robertsi (Kailola, 1990) - sedis mutabilis Potamosilurus velutinus (Weber, 1907) - sedis mutabilis Sciades Müller & Troschel, 1849 Sciades assimilis (Gùnther, 1864) Sciades bonillai (Miles, 1945) Sciades couma (Valenciennes, 1840) Sciades dowii (Gill, 1863) - sedis mutabilis Sciades emphysetus Müller & Troschel, 1849 Sciades felis (Linnaeus, 1766) Sciades guatemalensis (Günther, 1864) Sciades herzbergii (Bloch, 1794) Sciades leptaspis (Bleeker, 1862) Sciades mastersi (Ogilby, 1898) - sedis mutabilis Sciades passany (Valenciennes, 1840) Sciades paucus (Kailola, 2000) - sedis mutabilis Sciades platypogon (Günther, 1864) Sciades proops (Valenciennes, 1840) Sciades sagor (Hamilton, 1822) Sciades seemanni (Günther, 1864) Sciades sona (Hamilton, 1822) - sedis mutabilis Sciades utarus (Kailola, 1990) - sedis mutabilis Discussion and comparison with previous classifications The results obtained in this study are of primary importance for the systematics and taxonomy of the Ariidae for a long time a matter of controversy and misunderstanding among ichthyologists worldwide. The characterization of the genera, definition of their limits and species composition has been a great challenge within the systematics of the Siluriformes. Presently about 130 species are recognized as valid, but many need to be better characterized taxonomically. The difficulties in recognizing species identity and monophyletic taxa are mainly due to the wide geographic distribution of the group and the overall similarity in the external morphology of their representatives coupled with lack of adequate series of specimens in museum collections. Thus studies aimed at more comprehensive approaches on systematics and phylogeny of the Ariidae have not been entirely successful because of the above mentioned constraints. Recently Kailola (1990a, 2004), Betancur-R. & Mejía (2000), Betancur-R. (2003), Betancur-R. & Acero (2004) and Betancur-R. et al. (2004) based on the cladistic method, presented a preliminary analysis of the phylogenetic relationships of part of the genera and species of the Ariidae. Kailola (1990a) in an unpublished doctoral thesis discussed the relationships and zoogeography of the marine catfishes from New Guinea and Australia and Betancur-R. & Mejia (2000) and Betancur-R. (2003) did a similar study as part of the requirements for completion of the undergraduate program and a master’s dissertation respectively (not published) in Colombia and adjacent tropical waters. In both cases the authors examined only a geographically restricted subset of the recognized species and genera, limited to the areas above mentioned. Those studies included only a small number of species from African coasts, South America and Indian Ocean and did not redefine the complex genus Arius. The results obtained by Kailola (1990a) are essentially repeated in Kailola (2004) and the results obtained by Betancur-R. & Mejía (2000) are presented in Betancur-R. et al. (2004), but only part of the information contained in Betancur-R. (2003) is included in Betancur-R. & Acero (2004). In an unpublished Ph.D. study, Marceniuk (2003) included the largest number of ariid species ever assembled from different geographic areas of the world and used a vast array of morphological characters to study the systematics and phylogeny of the group. The present work is based on the results obtained by Marceniuk (2003). All nominal genera are revised and redefined through exclusive osteological characters and a combination of internal and external morphological characters. Based on examination of the type-species the following genera are considered valid: Arius, Aspistor, Bagre Cloquet 1816, Batrachocephalus, Brustiarius, Cathorops, Cephalocassis, Cinetodus, Cochlefelis, Cryptarius, Doiichthys, Galeichthys, Genidens, Hemiarius, Ketengus, Nedystoma, Nemapteryx, Neoarius, Netuma, Notarius, Osteogeneiosus, Pachyula and Sciades. The genera Plicofollis and Potamarius are considered valid through examination of species morphologically similar to the type-species. Representative material of Amissidens species-type was not examined and the genus is recognized exclusively on the basis of evidences presented by Kailola (2004). Amphiarius, Carlarius and Potamosilurus are described as new genera. The nominal genera Ailurichthys, Anemanotus, Ariopsis, Bagre Oken 1817, Felichthys, Glanis, Guiritinga, Hemipimelodus, Hexanematichthys, Pseudarius, Sciadeichthys, Sciadeops, Selenaspis, Septobranchus and Stearopterus are considered junior synonyms based on examination of the type-species and Leptarius and Pararius are considered junior synonyms based on examination of the type-species not cleared and stained. Ariodes and Tetranesodon are considered junior synonyms based on data presented by Kailola (2004). Breviceps Swainson, 1838 and Mystus Gray, 1854 are junior homonyms of names available for the genus-group and thus rejected. Catastoma and Sarcogeny