Cetopsis coecutiens ( Lichtenstein, 1819 )

Vari, Richard P., Ferraris Jr, Carl J. & de Pinna, Mário C. C., 2005, The Neotropical whale catfishes (Siluriformes: Cetopsidae: Cetopsinae), a revisionary study, Neotropical Ichthyology 3 (2), pp. 127-238 : 162-164

Cetopsis coecutiens ( Lichtenstein, 1819) Figs. 18 , 19 , Tables 9 -15

Silurus coecutiens Lichtenstein, 1819: 61 [type locality: Brazil]; 1823: 112 [ Brazil].– Cuvier, 1829: 292 [reference].

Cetopsis coecutiens .– Spix & Agassiz, 1829: 12, pl. 10, fig. 2 [rivers of equatorial Brazil].– Swainson, 1839: 308 [based on Spix &Agassiz, 1829].– Bleeker, 1858:258 [ Brazil]; 1862a: 403 [brief description]; 1862b: 16 [brief description, designation as type species of Cetopsis ]; 1863: 111 [brief description].– Cope, 1872: 292 [between mouth of the rio Negro and the Peruvian Amazon or Ucayale River (=río Ucayali)].– Eigenmann & Eigenmann, 1888: 157 [ Brazil: Gurupa]; 1890: 320 [redescription; lower Amazon basin]; 1891: 36 [in listing of Cetopsis species ].– Goeldi, 1898: 479 [Amazon].– Boulenger, 1897: 297 [ Brazil, Island of Marajo].– Regan, 1905: 190 [rio Negro].– Eigenmann, 1909: 322 [distribution]; 1910: 398 [distribution, not Irisanga citation].– Fowler, 1915: 228 [Ambiyacu River, andAmazon between mouth of rio Negro and Peru]; 1941: 472, fig. 30 [ Peru].– Schultz, 1944: 252 [literature records for Venezuela].– Fowler, 1945: 20, fig. 30 [ Peru].– Gosline, 1945: 54 [distribution, not Irisinga citation].– Van der Stigchel, 1947: 112 [ Brazil, Gurupá; redescription].– Fowler, 1954: 3 [literature compilation].– Dahl, 1960a: 306 [ Colombia, río Guayabero].– Ovchynnyk, 1967: 35 [ Ecuador, río Panayuca]; 1968: 255 [ Ecuador, río Panayuca].–Mago- Leccia, 1967: 255 [llanos of Venezuela]; 1970: 82 [ Venezuela, common name].– Cala, 1977: 11 [Colombian Orinoco; common name].– Ortega & Vari, 1986: 15 [Peruvian Amazon; common name].– Burgess, 1989: 292 [in listing of species].– Barriga, 1991: 56 [eastern Ecuador]; 1994a: 31 [ Ecuador, Parque Nacional Yasuni].–Lasso & Castroviejo, 1992: 74 [ Venezuela, río Guaritico; relative abundance].– Lasso et al., 1995: 18 [ Venezuela, río Guaritico; prey items].– Ferraris, 1996: 164 [form of adductor mandibulae muscle].– Taphorn et al., 1997: 85 [ Venezuela].– Chang, 1998: 26 [ Peru, río Madre de Dios basin, Tambopata-Candamo Reserved Zone].–Mojica-C., 1999: 565 [ Colombia, río Guaviare, río Meta].– Wallace, 2002: 312, fig. 125 [upper rio Negro; common name].–Evers & Seidel, 2002: 741 [listing].–Vari & Ferraris, 2003: 257 [in check list; synonymy, distribution, and common names].–[not Ringuelet & Arámburu, 1961: 46; Ringuelet et al., 1967: 348].

Cetopsis caecutiens .–Cuvier & Valenciennes, 1840: 384 [redescription].– Günther, 1864: 199 [ Brazil, rio Cupai].– Pellegrin, 1899: 158 [ Venezuela, río Apure].–Miranda- Ribeiro, 1912: 380 [ Brazil: Amazon to Cupay, rio Branco; redescription].– Starks, 1913: 34 [ Brazil: Pará and rio Madeira].– Miranda-Ribeiro, 1920: 15 [ Brazil, rio Madeira, mouth of rio Aripuanã].– Delsman, 1941: 80 [ Brazil, rio Trombetas and rio Óbidos].–[not Pozzi, 1945: 262; López et al., 2003: 37].

Cetopsis sp. – Goulding, 1981: 29, fig. 3.4 [ Brazil, rio Madeira, Teotônio rapids, feeding habits].–Castro &Arboleda, 1988: 12 [ Colombia, río Caqueta].– Ferreira et al., 1988: 346 [ Brazil, Roraima, rio Mucajai].– Goulding et al., 1988: 123 [ Brazil, rio Negro].– Lauzanne et al., 1991: 69 [ Bolivia, Trinidad].–Machado-Allison & Moreno, 1993: 86 [ Venezuela, río Orituco].

Cetopsis cf. caecutiens .– Mendes dos Santos et al., 1984: 78 [ Brazil, lower rio Tocantins; common names].

Diagnosis. Cetopsis coecutiens can be distinguished from all of its congeners by the combination of the presence of an eye, the conical teeth on the vomer and dentary, a body depth greater than 0.22 of SL, the elongated filaments on the distal portions of the first rays of the dorsal and pectoral fins in all specimens, and the presence of transversely-aligned, slit-like posterior nares.

Description. Body relatively stout, slightly-compressed anteriorly, becoming increasingly compressed posteriorly. Body depth at dorsal-fin origin approximately 0.23-0.27 of SL, and approximately equal to HL. Lateral line on body complete, unbranched, and midlateral; extending from vertical through pectoral-fin base onto hypural plate and terminating at, or slightly anterior to, posterior margin of hypural plate. Dorsal profile of body straight to slightly convex from nape to dorsal-fin origin, very slightly convex from dorsal-fin origin to caudal-fin base. Ventral profile of body convex along abdomen, approximately straight, but posterodorsally-slanted, along base of anal fin. Caudal-peduncle depth approximately equal to caudal-peduncle length.

Head triangular overall in lateral view and tapering to blunt point anteriorly. Dorsal profile of head slightly concave from tip of snout to nape. Ventral profile of head concave from margin of lower lip to slightly anterior of insertion of barbels and straight from that point to rear of isthmus. Snout obtusely pointed in dorsal view, with margins of head posterior to orbit slightly convex from dorsal view. Dorsal surface of postorbital part of head smooth, with minimal indication of underlying enlarged jaw musculature.

Opercular membrane attaching to isthmus posteriorly as far as vertical through pectoral-fin insertion. Opercular opening moderate; extending dorsal and ventral of pectoral-fin insertion by distance approximately equal to snout length.

Eye situated on lateral surface of head; located entirely dorsal to horizontal extending through pectoral-fin insertion; eye visible in dorsal view, but not in ventral view, of head. Middle of orbit located at approximately anterior one-fifth of HL. Eye diameter only slightly larger than width of anterior nares. Interorbital width approximately 0.40-0.42 of HL.Anterior narial opening circular, surrounded by very short, anteriorly-directed, tubular rim of skin. Opening of anterior nares located ventral of horizontal extending through tip of snout and approximately at horizontal running through maxillarybarbel origin. Distance between anterior nares equal to snout length. Posterior narial opening located on dorsal surface of snout and at horizontal extending through anterior margin of orbit; opening ovoid, with long axis of aperture aligned oblique to long axis of body. Anterior margin of posterior narial opening with low flap of skin. Distance between contralateral posterior nares only slightly larger than diameter of eye.

Mouth distinctly inferior; its width approximately equal to distance from tip of snout to posterior margin of orbit. Margin of lower jaw distinctly truncate, its posterior limit reaching to vertical through anterior margin of orbit. Premaxillary tooth patch elongate and continuous across midline; anterior margin convex and posterior margin concave. Premaxillary teeth relatively small, conical, sharply-pointed, and arranged in three irregular rows. Vomerine teeth very large, bluntly conical, and arranged in single, slightly-arched, transverse row. Dentary with single row of large, conical teeth approximately equal in number to, but 1.5 to 2.0 times size of, vomerine teeth.

Maxillary barbel very slender and short, its length approximately equal to distance from tip of snout to posterior margin of orbit; barbel origin located slightly anterior of vertical through anterior margin of orbit. Mental barbels approximately equal in size and length to maxillary barbel and to each other. Medial mental-barbel origin located just posterior of vertical through posterior margin of orbit. Lateral mental-barbel origin situated along vertical extending approximately one orbital diameter posterior of orbital margin. Tips of adpressed mental barbels falling distinctly short of posterior margin of opercle.

Dorsal fin relatively large overall with length of base of dorsal fin slightly less than one-half of HL. Length of longest branched dorsal-fin ray approximately two-thirds of HL. Dorsal-fin spinelet absent. First dorsal-fin ray not spinous, but with distal filament becoming progressively longer ontogenetically in both sexes; filament very well developed in largest specimens. Distal margin of dorsal fin slightly concave, with first ray longest. Dorsal-fin origin located at anterior one-third of SL and along vertical through middle of adpressed pectoral fin. Tip of adpressed dorsal fin, excluding distal filament on first fin ray, reaching to vertical through pelvic-fin insertion. Tip of distal filament of adpressed dorsal fin reaching to vertical through anal-fin origin. Posterior most dorsal-fin ray without posterior, membranous attachment to body.

Caudal fin deeply-forked, symmetrical; tips of lobes acutely pointed. Length of longest caudal-fin ray approximately two times length of middle fin rays.

Base of anal fin comparatively short, approximately onefourth of SL.Anal-fin origin located posterior of middle of TL. Anal-fin margin straight in females and immature males, with posterior most unbranched ray longest and lengths of subsequent rays becoming gradually shorter. Anal-fin margin in mature males slightly convex along anterior three-fourths of fin and then slightly concave along posterior five or six rays thereby giving appearance of slight lobe in anterior portion of fin. Posterior most anal-fin ray without posterior, membranous attachment to body.

Pelvic fin short, truncate, with distal margin nearly straight and first ray longest. Pelvic-fin insertion located approximately at middle of SL and entirely posterior to vertical through posterior terminus of base of dorsal fin. Tip of adpressed pelvic fin falling short of vent by distance approximately equal to length of first pelvic-fin ray. Medial most pelvic-fin ray with membranous attachment to body for basal one-half of its length.

Pectoral-fin length approximately two-thirds of HL. Pectoral-fin margin concave anteriorly and convex along posterior four rays. First pectoral-fin ray not spinous, but with distal filament; filament proportionally much longer in larger specimens. Tip of filament in adpressed dorsal fin reaching beyond vertical through pelvic-fin insertion in large individuals.

Coloration in alcohol. Head in medium to larger-sized specimens dusky dorsally with darker pigmentation becoming progressively diffuse ventrally. Smaller specimens with indistinct dusky band extending across interorbital region and with snout very pale. Body dusky dorsally, lighter ventrally, with relatively distinct ventral limit to field of dusky body pigmentation. Demarcation between dark and light pigmentation located dorsal to lateral line in examined specimens of all sizes.

Dorsal fin with dark pigmentation present on interradial membrane between first and second branched rays and with remainder of fin pale. Pectoral fin pale other than for dark pigmentation on interradial membrane between first and second rays. Pattern of darker pigmentation on dorsal and pectoral fins more pronounced in larger specimens. Caudal, anal, and pelvic fins without pronounced dark pigmentation.

Barbels pale to dusky.

Coloration in life. Overall coloration as in preserved specimens, but with light silvery-blue sheen on head and body.

Sexual dimorphism. The first rays of the dorsal- and pectoral-fin rays of both sexes of Cetopsis coecutiens have elongate, distal filaments that do not appear to demonstrate sexual dimorphism in relative length. The females and immature males have a straight anal-fin margin, whereas mature males have a largely convex fin, albeit with a slightly concave margin along its posterior portion (see description of form of anal fin above).

Distribution. Cetopsis coecutiens is broadly distributed through the Amazon basin in Bolivia, Brazil, Colombia, Peru, and Venezuela, and also inhabits the rio Tocantins basin in Brazil, and the río Orinoco basin in Colombia and Venezuela ( Fig. 19 ).

Common name. Brazil, rio Tocantins: “Candiru,” “Piracatinga” (Mendez dos Santos et al., 1984: 78), rio Madeira: “Candiruaçú ( Goulding, 1981: 31); Colombia:“Bagre ciego” ( Cala, 1977: 11); Peru:“Canero” ( Ortega & Vari, 1986: 15); Venezuela: “Bagre ciego” ( Mago-Leccia, 1970: 82).

Remarks. Lichtenstein (1819: 61) based his description of Silurus coecutiens on two specimens. One of the two syntypes of the species was listed in both the original catalogue of ZMB and also in the second catalogue of that museum (that was started around 1860) wherein it was cataloged as ZMB 3093 (H.-J. Paepke, ZMB; pers. commun., 1998). Recent searches through the ZMB collections failed to yield the syntype and that specimen is apparently no longer extant (H.-J. Paepke, ZMB; pers. commun., 1998). The second syntype was offered for sale by Lichtenstein (1823: 112) and there is no record of the disposition of that specimen.

The material that served as the basis of the description of Silurus coecutiens originated in Brazil ( Lichtenstein, 1823: 112). According to the ZMB catalogues the syntypes were collected by an individual named Sieber, whose collecting efforts were limited to the area of Belém, Pará, Brazil (H.-J. Paepke, ZMB; pers. commun., 1998), a region within the general known distributional range of Cetopsis coecutiens . Despite the apparent loss of both of the syntypes of the species, the distinctive features of Silurus coecutiens (the Cetopsis coecutiens of this study) detailed by Lichtenstein (1819: 61) make it clear that the name is appropriately applied to the material herein identified as that species.

The distributional range of Cetopsis coecutiens is by far the greatest of any species within the Cetopsinae and it is the only species in the subfamily with a broad distribution across both the rio Amazonas and río Orinoco basins. Despite this broad geographic distribution, detailed comparisons failed to reveal any meristic or coloration differences between the samples from those two drainage systems that would justify the subdivision of the C. coecutiens of this study into more than one species.

Mendez dos Santos et al. (1984: 78) cited Cetopsis cf. caecutiens from the lower portions of the rio Tocantins basin. Our results indicate that C. coecutiens is the only species that both occurs in that portion of the river basin and is likely to be cited as C. cf. caecutiens and we consider this citation to represent a record of the latter species.

Pozzi (1945: 262) followed by Ringuelet &Arámburu (1961: 46), Ringuelet et al. (1967: 348) and López et al. (2003: 37) cited Cetopsis coecutiens from northwestern Argentina The original Pozzi report (1945, map on page 251) indicated two localities in the Provinces of Salta and Jujuy from which species of the Cetopsinae originated, with both of those sites within the río Bermejo basin in the northwestern portion of the río de La Plata drainage system. None of the specimens of Cetopsis coecutiens examined in this study originated in the río de La Plata basin and the only species of the Cetopsinae that we examined that originated in the río Bermejo system was C. starnesi . Because of this evidence, we consider the citations of Cetopsis coecutiens from Argentina to be based on misidentifications of another species, most likely C. starnesi .

Material examined. 268 specimens (23-264 mm SL). Indefinite locality: Between the mouth of the rio Negro and the Peruvian Amazon or Ucayali River ,” ANSP 8384 , 1 (135) . Bolivia. Riberalta, lago Tumi Chacua , near río Beni (approximately 10°59’S, 66°06’W), ANSP 121606 , 1 (90) . Beni: río Mamoré, 5 km SW Buena Hora (= Boa Hora : 11 o 41’S, 65 o 05’W), Brazil, AMNH 55423 , 1 (127) . Río Mamoré , Trinidad (14 o 47’S, 64 o 47’W), MNHN 1988-1996 , 1 (193) . Brazil. Acre: Município de Artdira , rio Acre and marginal lagoons, LIRP 589 , 3 (128- 188) . Amapá: Fazendinha, rio Amazonas, near to Macapá (0 o 02’N, 51 o 03’W), MNRJ 16933 , 3 (57-99) . Amazonas: rio Madeira , AMNH 3870 , 1 (218) . At mouth of rio Putomayo (= rio Içá ; approximately 3 o 05’S, 67 o 57’W), CAS 65903, 1 (147) . Paraná de Janauaca (3 o 28’S, 60 o 17’W), USNM 217362 , 5 (175-195) ; MCZ 99231, 9 (166-264) . Lago Januacá (3 o 25’S, 61 o 21’W), MZUSP 24915 , 1 (139) . Rio Amazonas, Itacoatiara (3°08’S, 58°25’W), MHNG 2582.75 , 1 (195) . Fonte Boa, igarapé Jacaré, near Fonte Boa, MZUSP 23354 , 1 (113; specimen cleared and stained) . Mouth of lago Mamiá, below Coari (4 o 15’S, 63 o 03’W), MZUSP 23252 , 4 (132-168) . Rio Madeira, Borba (4 o 24’S, 59 o 35’W), MZUSP 28248 , 1 (154) . Rio Juruá, Carauari (4 o 52’S, 66 o 54’W), MZUSP 30697 , 1 (127) ; MZUSP 30692 , 1 (86) . Rio Madeira, Aripuanã (= Nova Aripuanã ) (5 o 06’S, 60 o 22’W), MZUSP 30695 , 1 (115) . Near Furo do Auozal , MZUSP 7401 , 1 (190) . Rio Içá at Cuiabá (2 o 57’S, 68 o 14’W), NRM 16534, 2 (128-160) . Rio Uaupés at Taracuá (= Taraquá ; 0 o 06’N, 68 o 28’W), NRM 17132, 1 (173) ; NRM 25551, 1 (195) ; NRM 25552, 1 (218) . Rio Uaupés at Jauareté (= Iaureté ) (0 o 36’N, 69 o 12’W), NRM 25554, 1 (183) . Matupiri (5 o 33’S, 61 o 15’W), NRM 23948, 1 (62) . Rio Negro drainage (locality information questionable), NRM 25555, 1 (190) . Rio Negro, 10.4 km below rio Demimi (1°13’01"S, 62°14’29"W), MZUSP 55687 , 1 (37) . Rio Negro, 6.5 km below Jufari (1°17’37"S 61°56’57"W), MZUSP 55647 , 8 (48-78) . Rio Negro, near Baia do Cuieirus , MZUSP 38765 , 6 (2 specimens, 107-128 mm, cleared and stained) . Rio Purus , 24.3 km below paraná de Ananás, MZUSP 55697 , 4 (122-163) . Rio Solimões, 60 km above paraná do Barrosa (3°11’59"S, 59°54’27"W), MZUSP 57770 , 2 (31-76) . Rio Solimões, lago above Purus (3°49’56"S, 61°39’05"W), MZUSP 57340 , 1 (46) . Rio Purus , Caitaú, MZUSP 30443 , 1 (97) . Rio Solimões , 25 km below paraná de Amaturá (3°10’48"S, 67°55’40"W), MZUSP 55682 , 1 (28) . Rio Solimões , 20 km below Içá (3°00’45"S, 67°52’42"W), MZUSP 55684 , 1 (25) . Rio Solimões, 15 km below paraná Iranduba (3°14’22"S, 59°53’53"W), MZUSP 55646 , 2 (23-30) . Goiás: rio Tocantins between Porto de Garimpo and Foz do rio Preto , MHNG uncataloged (field number BR 954 ), 1 (215). Rio Tocantins between site of future dam of Usina Hidroeléctrica Cana Brava and Ilha, MNRJ 12907 , 1 (198) . Município de Minaçu / Cavalcante, rio Tocantins , at site of future dam of Usina Hidroeléctrica Serra da Mesa (13 o 50’S, 48 o 19’W), MNRJ 13235 , 2 (200-264) . Rio Vermelho, above Aruanã (approximately 14 o 54’S, 51 o 05’W), MZUSP 38826 , 1 (220) . Rio Araguaia, Aruanã (approximately 14 o 54’S, 51 o 05’W), MZUSP 4863 , 1 (240) . Matto Grosso: rio Araguaia , lago Dumbá Grande, MZUSP 24819 , 3 (168-197) . Rio Araguaia , lago Montario, MZUSP 54176 , 1 (202) . Rio Araguaia , near Ilha do Biratã, MZUSP 62518 , 1 (162) . Pará: no specific locality, CAS-SU 22197 , 1 (182) . Rio Tocantins, in front of Tucuruí (3 o 42’S, 49 o 27’W), MZUSP 24177 , 2 (183, other specimen cleared and stained and disarticulated) . Rio Tocantins , from immediately below to 3 km below Tucuruí reservoir (3°42’S, 49°27’W), MZUSP 61996 , 1 (179) . Rio Xingu, Belo Monte (3°10’S, 51°50’W), MZUSP 59092 , 4 (173-237) . Rio Amazonas, Maiacuru (2 o 14’S, 54 o 17’W), MNRJ 9395 , 15 (73-120) . Santarém (2 o 26’S, 54 o 42’W), FMNH 69823 , 1 (191) . Rio Trombetas, mouth of lago Jacaré, Reserva Biologica de Trombetas , MZUSP 15511 , 1 (170) . Mouth of lago Jacaré, Reserva Biologica de Trombetas , MZUSP 15533-34 , 2 (177-180) . Praia do lago near mouth of lago Jacaré, Reserva Biologica de rio Trombetas , MZUSP 15826 , 1 (195) . Rio Trombetas, Oriximiná (1 o 45’S, 55 o 52’W), MZUSP 5466 , 1 (170) . Rio Trombetas, mouth of lago Paru (1 o 57’S, 55 o 52’W), MZUSP 5685 , 1 (97) . Rio Tapajós, Ilha Tapaiúna (2 o 54’S, 55 o 05’W), MZUSP 21985 , 5 (179-192) . Santana, Furo dos Macacos, MZUSP 23653 , 1 (172) . Abaetetuba, rio Tocantins (1 o 42’S, 48 o 54’W), MZUSP 23945 , 3 (109-130) . Rio Tapajós , Maloquinha, MZUSP 24308 , 2 (157-162) . Rio Amazonas, Furo Ituquara below Gurupá (approximately 1 o 25’S, 51 o 39’W), MZUSP 43228 , 12 (97-153) ; MZUSP 43288 , 4 (143- 186) . Ilha Urutaí, Município de Gurupá (approximately 1 o 25’S, 51 o 39’W), MZUSP 9419 , 1 (146) . Rio Pará, Ilha do Camaleão (approximately 1 o 30’S, 48 o 05’W), MZUSP 9474 , 4 (113-153) . Corcorado (1 o 51’S, 49 o 02’W), MZUSP 9475 , 2 (144-158) . Rio Amazonas, mouth of Furo do Aququi (1 o 35’S, 52 o 35’W), MZUSP 9491 , 2 (107-123) . Probably in vicinity of Belém, NRM 7102 , 1 (137) . Engenho Santo Antonio, Furo de Panaquera , MZUSP 23975 , 8 (143-165; 1 specimen, 145 mm, cleared and stained) ; ROM 37970, 3 (134-142) . Rondônia: rio Madeira, Foz do Jamarí (8 o 45’S, 63 o 27’W), MZUSP 28370 , 11 (154-205) . Tocantins: rio Tocantins at Porto Nacional (10 o 42’S, 48 o 25’W), CAS 6609 , 1 (218) . Rio Tocantins, Município de Peixoto , MZUSP 45102 , 2 (136-145) . Colombia. Amazonas: Leticia, río Amazonas , ANSP 119877 , 1 (93) . Caqueta: río Orteguaza , MHNG 2526.66 , 22 (106-120) . Río Amazonas , Isla de Santa Sofia II, approximately 20 mi (= 32 km) NW of Leticia, CAS 40714, 1 (117) . Peru. Amazonas: río Santiago , 1.5 km upstream of La Poza (latter locality at 4 o 01’S, 77 o 45’W), LACM 39864-1 , 1 (197) . Río Marañon, Bagua, Ciro-Cenepa (4 o 42’S, 78 o 10’W), MUSM 12494 , 1 (195) . Loreto: río Ambiyacu , ANSP 21533 , 1 (135; originally cited in literature as from Ecuador) . Río Ambiyacu, near Pebas (approximately 3 o 20’S, 71 o 49’W), CAS SU 34225 , 1 (101) ; CAS SU 36191 , 1 (38) . Rio Amazonas , INHS 38932 , 1 (118) . Río Napo, Puesto de Vigilancia Arcadia (0 o 59’S, 75 o 18’W), MUSM 6023 , 1 (210) . Río Aguarico, Puesto de Vigilancia Castana (0 o 48’S, 75 o 14’W), MUSM 6024 , 1 (220) . Madre de Dios: río Madre de Dios , Albergue Cuzco-Amazonico, CAS 59457, 2 (110-119) . Río Madre de Dios , 2 km W of Puerto Maldonado (approximately 12 o 36’S, 69 o 11’W), CAS 78929, 2 (121-176) . Ucayali: río Ucayali, Pucallpa (8 o 23’S, 74 o 32’W), MZUSP 26418 , 1 (100) . Venezuela. Río Orinoco, Puerto Cabrian to Puerto Morocota, 149-150 nautical mi (=276-277.8 km) from sea buoy, ANSP 152873 , 1 (68) . Amazonas: río Negro at San Carlos de río Negro (1 o 58’N, 67 o 04’W), AMNH 74450 , 1 (185) . Anzoátegui: Caño tributary to río La Pena, left bank tributary of río Orinoco , opposite Ciudad Bolivar (approximately 8°10’N, 63°33’W), ANSP 152872 , 1 (31) . Río Orinoco at Soledad boat launch just below Puente Angostura (approximately 8°10’N, 63°34’W), ANSP 160997 , 1 (51) . Apure: río Apure, between mouth of río Portuguesa and San Fernando de Apure airport, ANSP 165235 , 1 (137) . Río Apure, above Apurito (7 o 56’20"N, 68 o 28’50"W), MCNG 11234 , 1 (123) . Río Sarare , between Las Bocas and Guasdualito (7 o 11’50"N, 71 o 48’10"W), MCNG 5262 , 2 (183-216) . Río Apure, just W of Apurito, on S bank (7 o 56’N, 68 o 27’W), TNHC 13855 , 2 (199-210) . Delta Amacuro: río Orinoco , between Puerto Ordaz and Barrancas, AMNH 58156 , 3 (56-104) ; AMNH 58156 SW, 1 (24.6, cleared and stained). Río Orinoco, near downstream end of Isla Muasimoina , 44 nautical mi (=81.5 km) from sea buoy (8 o 36’36"N, 60 o 45’W), USNM 265631 , 1 (157) . Río Orinoco channel, N of Isla Tres Caños , 128 nautical mi (=237.1 km) from sea buoy (8 o 39’N, 61 o 58’W), USNM 265660 , 1 (28) . Río Orinoco channel, at Puerto Ordaz , N of Islote Fajardo, 180 nautical mi (=333.4 km) from sea buoy (8 o 23’33"N, 62 o 39’54"W), USNM 265673 , 1 (34) ; LACM 43102-3 , 2 (15-21, larger specimen cleared and stained) . Río Orinoco tributary across from Isla Tres Caños , on N side of river, 131.8 nautical mi (=224.1 km) from sea buoy (8 o 39’48"N, 62 o 01’W), USNM 265707 , 1 (19) . Río Orinoco , in deep channel (8°23’N, 61°02’W), CAS 58079, 6 (102-170) . Río Orinoco , in deep channel (8°22’N, 62°40’W), CAS 58161, 1 (33) . Guarico: río Orinoco, at Laguna Isla Isabella , cross outlet of lagoon, 201 nautical mi (=372.3 km) from sea buoy (8 o 18’N, 63 o 56’W), USNM 265628 , 30 (112-225; 1 specimen cleared and stained) . Río Orinoco channel, between Puerto Ordaz and Ciudad Bolivar, 202 nautical mi (=374.1 km) from sea buoy (8 o 18’N, 63 o 56’W), USNM 265657 , 2 (37-54) . Portuguesa: río Portuguesa (9 o 04’00"N, 69 o 30’00"W), MCNG 15775 , 3 (185-192).

Figures

Fig. 18. Cetopsis coecutiens, MCNG 5262, 216 mm SL; Venezuela, Apure, río Sarare, between Las Bocas and Guasdualito (7o11’50"N, 71o48’10"W).

Fig. 19. Map of central and northern SouthAmerica showing geographic distribution of Cetopsis coecutiens (type locality inexact = Brazil), Cetopsis fimbriata (1 = holotype locality), and Cetopsis gobioides (2 = lectotype locality of Cetopsis chalmersi) (some symbols represent more that one locality and/or lot of specimens).