Salminus iquitensis ( Nakashima, 1941 ) Lima, 2022

Lima, Flávio C. T., 2022, Revision of the smaller-sized dorados (Salminus), with comments on the monophyly of the genus and its biogeography (Characiformes: Bryconidae), Zootaxa 5226 (1), pp. 1-66 : 32-44

Salminus iquitensis ( Nakashima, 1941) , new combination

( Figs. 12–13 )

Salminus hilarii (not Valenciennes): Pellegrin, 1899: 157 ( Venezuela, río Apure); Pearson, 1924: 49 (Ixiamas, río Beni basin, Bolivia); La Monte, 1935: 8 (“Rio Macauhan, tributary of Rio Yaco (...) tributary of rio Purus, 69’W, 9°20’S ” = Brazil, Acre, rio Macau ã, trib. rio Iaco); Fowler, 1943: 243–244 ( Colombia, Florencia, río Orteguaza); Mago-Leccia, 1970: 73 (listed, Venezuela; common names); Saul, 1975: 110 (río Conejo, Napo, Ecuador); Ortega & Vari, 1986: 9 (listed, Peru); Ferreira et al., 1988: 344 ( Brazil, Roraima, rio Mucajaí, Ilha Pared ã o); Géry & Lauzanne, 1990: 120, 122 (in part; río Apure, Venezuela; diagnosis in key); Barriga, 1991: 27 ( Ecuador, cisandean basins; common name); Lasso, 1992: 13 (lower río Suapure, río Orinoco basin, Venezuela); Taphorn, 1992: 321–323 ( Venezuela, río Apure basin; habitat, diet, migrations, common names); Winemiller et al., 1996: 23, 32 (río Orinoco basin, Venezuela: migrations, conservation); Taphorn et al., 1997: 74 (listed, Venezuela); Salinas-Coy & Agudelo-Córdoba, 2000: 57–58 (río Putumayo, río Caquetá and río Guaviare, Colombia; fisheries); Silvano et al., 2001: 60–61 (photo; Acre, upper rio Juruá; common name); Diaz-Sarmiento-Diaz & Alvarez-Léon, 2003: 313 (río Caquetá, Colombia; fishery); Lima et al., 2003: 156 (in part: Holobrycon iquitensis as a synonym; distribution, common names); Amaral, 2004: 80 (upper rio Juruá basin, Acre, Brazil; common name, importance in fisheries); Wright & Flecker, 2004: 447, 449 (río Las Marias, río Apure basin, Venezuela); Lasso, 2004: 156–158 (Estación Biológica El Frío, caño Guaritico, Apure, Venezuela; occurrence, diet, habitat); Bogotá-Gregory & Maldonado-Ocampo, 2006: 69 (listed, Colombia, Amazon basin); Ortega et al., 2006: 103 (río Putumayo, Colombia / Peru; listed); MaldonadoOcampo et al., 2008: 173 (listed, Colombia, Amazon and Orinoco basins); Morales-Betancourt & Lasso, 2011: 253–254 (photo, short description, distribution, habitat, reproduction, migrations, fisheries; Amazon and Orinoco basins, Colombia); Ortega et al., 2011: 37 (listed, Peru); Barriga, 2012: 108 ( Ecuador, río Napo basin; occurrence); Usma-Oviedo et al., 2013: 290–291 (Amazon and Orinoco basins, Colombia; photo, common names, short description, distribution, habitat, reproduction, migrations, fisheries); Dagosta & de Pinna, 2019: 89 (in part; “Guaporé, Beni-Madre de Dios, Putumayo, Japurá, Branco”); Meza-Vargas et al., 2021: 20 (listed, Peru, Departamento Loreto); Zamudio López, 2022: 91 ( Colombia, Arauca, río Cinaruco; photo in life).

Salminus affinis (not Steindachner): Eigenmann, 1922: 158 (in part; río Chanchamayo, La Merced, Peru); Eigenmann & Allen, 1942: 260 (río Ucayali, Orellana, Peru); Ortega et al., 1977: 14, 37, pl. IIC (río Ucayali, Pucallpa, Peru); Ortega & Vari, 1986: 9 (listed, Peru); Géry & Lauzanne, 1990: 123 (in part; diagnosis in key); Barriga, 1991: 27 ( Ecuador, cisandean basins; common name); Ortega, 1996: 467 (río Manu, Parque Nacional Manú, Depto. Madre de Dios / Cuzco, Peru); Chang, 1998: 23 (río Tambopata basin, Zona Reservada Tambopata-Candamo, Deptos. Madre de Dios / Puno, Peru); Chernoff et al., 2000: 280 ( Bolivia, Amazon basin); Goulding et al., 2003: 137–138 (photo; río los Amigos, Depto. Madre de Dios, Peru); Diaz-Sarmiento & Alvarez-Léon, 2003: 313 (río Caquetá, Colombia; fishery); Bogotá-Gregory & Maldonado-Ocampo, 2006: 69 (listed, Colombia, Amazon basin); Renjifo, 2007: 198, 200 (upper río Yuruá, Depto. Ucayali, Peru; occurrence, common names); Maldonado-Ocampo et al., 2008: 173 (in part; listed, Colombia, Amazon basin); Jiménez-Segura et al., 2011: 249–250 (in part: río Putumayo and río Caquetá basins, Colombia; based on Bogotá-Gregory & Maldonado-Ocampo, 2006); Ortega et al., 2011: 37 (listed, Peru); Usma-Oviedo et al., 2016: 113 ( Colombia, río Meta; occurrence); DoNascimiento et al., 2017: 63 (in part; listed, Colombia, Amazon and Orinoco basins); Quezada García et al., 2017: 337 ( Peru, Ucayali, río Aguaytía basin; occurrence); Lasso et al., 2021: 150–152 ( Colombia, río Orinoco basin; diagnosis, photo in life, distribution, natural history, recreational fisheries); Meza-Vargas et al., 2021: 20 (listed, Peru, Departamento Loreto).

Holobrycon iquitensis Nakashima, 1941: 72–73 , fig. (type locality: “Cercanias del Puerto de Iquitos”, Peru); Fowler, 1950: 305 (as a synonym of Salminus maxillosus ); Géry & Lauzanne, 1990: 116 (as a presumable synonym of Salminus hilarii ); Lima et al., 2003: 156 (as a synonym of Salminus hilarii ).

Salminus maxillosus (not Valenciennes): Fowler, 1945: 105 (Morelia, río Caquetá, Colombia); Ortega & Vari, 1986: 9 (listed, Peru); Barriga, 1991: 27 ( Ecuador, cisandean basins; common names); Diaz-Sarmiento & Alvarez-Léon, 2003: 313 (río Caquetá, Colombia; fishery).

Brycon iquitiensis: Howes, 1982: 32 (as a presumable synonym of Salminus hilarii ).

Salminus brevidens (not Cuvier): Barriga, 1991: 27 ( Ecuador, cisandean basins; common name).

Salminus cf. hilarii (not Valenciennes): Stewart et al., 1987: 29 (río Napo, Ecuador); Galacatos et al., 2004: 49 (Río Yasuni, Ecuador).

Salminus sp. : Lasso, 2001: 239 (río Negro, tributary of río Itenez, Depto. Santa Cruz, Bolivia); Galvis et al., 2006: 231, pl. 46 ( Colombia, Amazonas, Leticia; occurrence); Rodríguez-Olarte & Taphorn, 2006: 73–79 (río Orinoco basin, Venezuela; natural history); Carvalho et al., 2012: 976, 987 (río Las Piedras, río Madre de Dios basin, Madre de Dios, Peru; occurrence, photo); Jégu et al., 2013: 127 (río Iténez/Guaporé basin, Bolivia / Brazil; occurrence; based in Lasso, 2001); Lima et al., 2013: 344–345 (short description, photo; middle rio Madeira basin, Rondônia, Brazil); Urbano-Bonilla et al., 2018: 86 (río Cusiana, río Meta basin, Colombia; occurrence); Silva et al., 2020: 5, 8 (fig. 6) ( Brazil, Acre, rio Acre; photo).

Salminus brasiliensis (not Cuvier): Barriga, 2012: 108 ( Ecuador, río Napo basin; occurrence).

Diagnosis. Salminus iquitensis can be diagnosed from all congeners, except S. santosi , by the presence of numerous, tiny stripes concentrated at the interradial membranes of middle and central portions of caudal fin (vs. absence of such stripes in S. affinis , S. brasiliensis , S. franciscanus , and S. hilarii ). Salminus iquitensis can be additionaly diagnosed from all congeners, except S. affinis and S. santosi , by the presence of a dark, straight postorbital stripe extending across the contact area between infraorbitals 4 and 5 to upper portion of opercle (vs. absence of postorbital stripe S. brasiliensis , S. hilarii , and S. franciscanus ). Salminus iquitensis can be additionally diagnosed from S. brasiliensis by presenting considerably lower scale counts, i.e., lateral line scales (65–87, modally 75, vs. 79–102, modally 96), and horizontal scales between dorsal-fin origin and lateral line (11–14, modally 12, vs. 14–18, modally 16), and by possessing color in life mainly silvery, with golden pigmentation restricted to facial bones and underside (vs. overall color golden in living specimens). Salminus iquitensis can be diagnosed from S. santosi by possessing higher scale counts, i.e., lateral line scales (65–87, modally 75, vs. 61–74, modally 69), and horizontal scales between dorsal-fin origin and lateral line (11–14, modally 12, vs. 10–13, modally 11), higher anal-fin branched rays counts (20–25, modally 23, vs. 18–24, modally 22), and a distinct color pattern of the caudal fin in living specimens, with distal portion typically rosy or light-red (vs. distal portion bright red). See the item “Remarks” of S. santosi for additional comments regarding the diagnosis of both species.

Description. Morphometric data presented in Table 4–5 . Middle-sized species, larger specimen examined 382.0 mm SL. Body elongated, largest body height at level of dorsal-fin origin. Dorsal profile slightly convex from snout tip to vertical through anterior nostril, straight to slightly concave from later point to tip of supraoccipital process, slightly convex from later point to dorsal-fin origin, straight along dorsal-fin basis, and slightly convex from dorsalfin terminus to adipose-fin origin. Dorsal profile of caudal peduncle slightly concave. Ventral profile slightly convex from tip of lower jaw to pelvic-fin insertion, straight to slightly convex from later point to anal-fin origin, and approximately straight along anal-fin basis. Ventral profile of caudal peduncle slightly concave.

Head profile acute anteriorly (often more so in specimens from río Orinoco basin), mouth terminal. Maxillary elongated, extending posteriorly slightly beyond vertical through posterior eye margin, extending even slightly beyond in some specimens from río Orinoco basin (MCNG 1723, MCNG 35450). Adipose eyelid well developed. Premaxilla with two teeth rows. Teeth of outer row teeth considerably larger than those of inner series other than for second tooth of inner row. Outer row with 6–11 teeth, approximately equal in size; teeth with distinct, elongate basal portion (shaft), and apical portion (crown). Crown triangular, angles possibly constituting poorly differentiated cusps. Inner tooth row with 7–17 teeth, symphyseal tooth relatively large, second tooth slightly smaller, third and subsequent teeth considerably smaller. Teeth of inner row morphologically similar to those from outer row, except for proportionally shorter shafts and being more massive overall. Middle portion of ventral margin of maxilla slightly concave. Maxilla with 23–47 teeth, similar in shape to those from outer row of premaxilla, but slightly smaller and decreasing very gradually in size posteriorly, with less developed crowns, last teeth roughly conical. Dentary with 18–37 teeth on primary, outer row morphologically similar to those from outer row in premaxilla, and other than first to third teeth, slightly smaller than those on premaxilla. Remaining teeth gradually decreasing in size and presenting less developed crowns. Teeth of outer dentary row, outer premaxilla row, and maxilla with crowns slightly recurved lingually. Inner tooth row with 38–64 conical teeth arranged continuously from symphysis to terminus of inner rim of replacement teeth trench. Teeth at inner row oriented at right angle relative to teeth of primary row, with apices directed lingually.

Scales cycloid. Lateral line complete, extending from supracleithrum to caudal-fin base. Lateral line scales 65(1), 66(2), 67(4), 68(4), 69(10), 70(5), 71(13), 72(12), 73(15), 74(8), 75(17), 76(6), 77*(7), 78(9), 79(7), 80(1), 81(4), 82(1), or 87(1). Laterosensory tubes simple, straight or deflected downward. Horizontal scale series between dorsal-fin origin and lateral line 11(11), 12*(65), 13(44), or 14(7). Horizontal scale series between lateral line and pelvic fin insertion 4(3), 5(19), 6*(73), 7(29), or 8(3). Circumpeduncular scales 21(6), 22(17), 23(35), 24(40), 25*(19), 26(2), 27(6), or 28(2).

Dorsal-fin rays ii, 9. Dorsal-fin origin approximately midway between snout and hypural joint. First dorsal-fin pterygiophore inserting behind neural spine of 14 th (15) or 15 th (2) vertebrae. Anal fin rays iv, 20(3), 21(14), 22(24), 23*(48), 24(22), or 25(8). First anal-fin pterygiophore inserting behind hemal spine of 25 th (10), 26 th (5), or 27 th (1) vertebrae. Last unbranched rays and first to second branched anal-fin rays longer; third to ninth ray gradually shortening, with remaining rays approximately equal in size. Pectoral fin-rays i, 12(7), 13(42), 14*(60), or 15(11). Pelvic-fin rays i, 7, a single specimen i, 6. Principal caudal-fin rays 10/9. Anal fin with small hooks along last unbranched ray and anteriormost branched anal-fin rays in seven specimens (IAvH-P 1940, 230.9 mm SL; IAvH-P 2906, 245.0 mm SL; MUSM 29, 2, 235.8– 250.6 mm SL, MZUSP 26078, 250 mm SL; USNM 229054, 179.6 mm SL; MPUJ 2943, 215.2 mm SL). Hooks limited to posterior branch of ray. Pelvic fins with hooks on branched rays 1–6 (MZUSP 26708, MPUJ 2943), limited to posterior branch of ray. Scales sheath composed of two horizontal series covering basal portion of anal-fin rays. Caudal fin moderately forked to slightly emarginated. Laterosensory tube on caudal fin extending to caudal-fin terminus, with dorsally and ventrally oriented side branches along its length. Central caudal-fin extension present, discernible in specimens with well-preserved caudal-fin margins, small in most specimens (e.g., Fig. 13B ), relatively well-developed in some specimens (e.g., Fig. 13D ), at level of main rays 10–11, where laterosensory tube ends.

Four branchiostegal rays. First gill arch with 15(2), 16(1), 17(1) lower gill-rakers, 15(1) or 16(3) upper gill rakers and 1 (4) at angle. Vertebrae 46(2), 47(5), 48(7), or 49(2). Supraneurals 11(12), or 12(3).

Color in alcohol. Overall color of specimens still retaining guanine grey dorsally, with plumbeous tint. Top of head (including supraorbital), snout and anterior portion of maxilla brown; remaining infraorbitals and opercle silvery. Maxilla, gular area and dentary light brown. Dark postorbital stripe approximately straight, extending across contact area between infraorbitals 4–5 to upper portion of opercle, formed by concentration of small dark chromatophores, variously developed but always present. Sides of body clear, with silvery hue, dark grey dorsally. Humeral blotch present, little conspicuous, oval-shaped, horizontally elongated, formed by pigmentation subjacent to scales, lying immediately above lateral line, at level of second to sixth scales. Narrow, straight dark stripes extending along trunk, formed by dark chromatophores concentrated at central-distal portion of each scale. Stripes present across all trunk, more conspicuous dorsally. Broad dark stripe across caudal peduncle, starting at vertical through immediately behind adipose fin, and gradually broadening towards caudal-peduncle terminus. Caudal-peduncle stripe extending into distal portion of four middle caudal-fin rays. Caudal fin with numerous, tiny stripes concentrated at the interradial membranes of middle and central portions of caudal fin, formed by rows of dark chromatophores. Anal, dorsal, pelvics, pectorals, and adipose fins clear, with few, scattered dark chromatophores. Specimens which lost guanine pigmentation as a result of long storage in formalin with overall color brown, lacking silvery pigmentation on sides of body, infraorbitals, or opercle.

Color in life. Description based on pictures of specimens from the río Meta (río Orinoco basin) and from the río Putumayo (río Amazonas basin) in Colombia by A. Linares, from a specimen from a tributary of río Guayabero (CZUT-IC 7147) by A. Urbano-Bonilla, from subaquatic pictures of specimens taken at the rio Guejar and Caño Sardinata (río Guaviare basin, Colombia) by O. Lucanus, from the río Aguaytia (río Amazonas basin, Peru) by A. Linares and M. Hidalgo, from a tributary of the río Inambari (río Madre de Dios basin, Peru), by J.M. Rodriguez Vera, from a freshly collected specimen from the rio Abun ã, rio Madeira drainage (MCP 37564) taken by the author, from a specimen (now lost) collected from a tributary of the rio Branco near Serra Grande (Cantá), Roraima, by F.A.G. Melo, from three specimens from rio Acari, rio Branco basin, provided by Í. Mesquita and S.R. Briglia-Ferreira (two of them preserved: ZUEC 17546), from a small specimen from the río Los Amigos, río Madre de Dios drainage ( Goulding et al., 2003: 138; presumably MUSM 19149), from a specimen from the río Las Piedras, Peru ( Carvalho et al., 2012: 987), and from a specimen lacking a specific locality (Lasso et al., 2011: 251). Sides of body clear, with silvery hue; dorsum grey. Golden-yellow pigmentation on facial bones and opercle (silvery in small specimens). Eye rim light-yellow to olive. Caudal fin light yellow at basis, rosy to light red at distal margin (deep red in some specimens, e.g., Lasso et al., 2011: 251; Carvalho et al., 2012: 987). Dorsal, anal, pectorals, pelvics, and adipose fin yellowish to orangish. Dark markings as in preserved specimens, except that some scales with orangish to reddish pigmentation, forming broken, small vertical stripes ( Fig. 13 ).

Sexual dimorphism. As described in the Description, fin hooks are present at pelvic and anal fins of mature males (IAvH-P 1940, 230.9 mm SL; IAvH-P 2906, 245.0 mm SL; MUSM 29, 2, 235.8– 250.6 mm SL, MZUSP 26078, 250 mm SL; USNM 229054, 179.6 mm SL).

Geographical variation. No meaningful morphometric, meristic, or color pattern differences were found between the populations of S. iquitensis from the western Amazon and the río Orinoco basins ( Tables 5–7 ). We have tentatively assigned the population of the rio Branco basin to S. iquitensis based on the fact that the few specimens available from that basin (four specimens: INPA 10791, 1; NMW 56870, 1; ZUEC 17546, 2) have meristic features more compatible with this species than with S. santosi from the rio Tocantins basin (see Tables 6–7 ), and additionally because the color pattern of living specimens is more similar to specimens of S. iquitensis than to S. santosi . Machado et al. (2017, 2018) have not sequenced any specimens from the populations of the río Orinoco and rio Branco basins, so it is presently unknown whether there is any molecular divergence among the three disjunct populations herein assigned to S. iquitensis . See comments regarding the diagnosis between S. iquitensis and S. santosi on the item “Remarks” of S. santosi .

Common names: Brazil: “tabarana”, “tubarana” ( Silvano et al., 2001; S.R. Briglia-Ferreira, pers. comm.), “sh ã w ã w ã” (Kaxinawá language; Amaral, 2004); Colombia: “choja”, “pez lagartija” (Salinas-Coy Córdoba-Agudelo, 2000), “dorado”, “salmón” ( Diaz-Sarmiento & Alvarez-Léon, 2003), “saltarín”, “picuda”, “chojo”, “saltador” ( Usma-Oviedo et al., 2013); “picuda-llanera” ( Lasso et al., 2021); Peru: “sábalo”, “sábalo macho” ( Nakashima, 1941; Ortega & Vari, 1986); “sabalito” ( Goulding et al., 2003); “ypanha” (Amahuaca language; Renjifo, 2007); “sani” (Yaminahua language; Renjifo, 2007); Ecuador: “dama”, “damita” ( Barriga, 1991); Venezuela: “dorado”, “dorada”, “saltador”, “saltadora”, “sauta” ( Mago-Leccia, 1970; Lasso, 1992; Taphorn, 1992).

Distribution. Salminus iquitensis has a wide, though disjunct, distribution across the upper Amazon basin, from the upper rio Madeira basin to the río Caquetá basin, in Bolivia, Brazil, Peru, Ecuador, and Colombia, at the río Orinoco basin in Colombia and Venezuela, and at the rio Branco basin in Brazil ( Fig. 14 ). There are few records for the species from the rio Branco basin, and Fig. 14 includes information of one locality for the species from which the voucher specimen was unfortunately lost (a tributary of the rio Branco near Serra Grande, Cantá; F.A.G. Melo and S.R. Briglia-Ferreira, pers. comm.). A specimen supposedly collected at the río Pamplonita, a tributary of the Lago Maracaibo in Colombia (USNM 100771) by Brother Níceforo Maria in 1935 is herein considered to have incorrect locality data as there are no other records for Salminus for any rivers draining into the Lago de Maracaibo basin (e.g., Rodriguez-Olarte et al., 2009).

Ecological notes, conservation. Salminus iquitensis occurs mostly at piedmont localities at the río Orinoco, being uncommon in the lowlands ( Taphorn, 1992; Rodríguez-Olarte & Taphorn, 2006). The same is true for the Amazon basin, where the species clearly demonstrates a preference for piedmont areas, being rarely found at altitudes below 200 meters a.s.l. Records below this altitude (e.g., at the lower río Napo and other tributaries of the río Amazonas near Iquitos) are scarce, judging at least from the number of specimens collected at this relatively wellsampled area when compared with the amount of specimens available from less-intensively sampled areas near or at piedmont areas as the middle río Ucayali basin near Pucallpa, Peru. Rodríguez-Olarte & Taphorn (2006) published a detailed study on the natural history of S. iquitensis (identified by them as Salminus sp. ) from the piedmont rivers of the Orinoco basin in Venezuela, and most of the information below is summarized from that paper. The species is reported to favor rapids and fast-flowing river stretches (e.g., Saul, 1975). At the Orinoco andean piedmont area, small specimens (<20 cm TL) were observed in mixed schools with Brycon whitei , while larger specimens form small monospecific schools ( Rodríguez-Olarte & Taphorn, 2006). Females grow larger than males, and the smaller mature female reached 19.5 cm SL ( Rodríguez-Olarte & Taphorn, 2006). The smallest mature male examined (judging for the presence of hooks on the anal- and pelvic-fins) reached 179.6 mm SL (USNM 229054). Female fecundity ranged between 17,528 and 57,834 oocytes, and maximum ripeness was observed in June, with post-reproductive individuals of both sexes being found in July and August ( Rodríguez-Olarte & Taphorn, 2006). The species is reported to undertake reproductive migrations between piedmont areas and lowlands ( Taphorn, 1992; Winemiller et al., 1996; Rodríguez-Olarte & Taphorn, 2006; Usma-Oviedo et al., 2013). Salminus iquitensis was observed to be mainly active at early morning and late afternoon ( Rodríguez-Olarte & Taphorn, 2006). The species feeds primarily on small characins, with allochtonous invertebrates being the second most important feeding item (RodríguezOlarte & Taphorn, 2006). Large specimens (> 20 cm SL) at the Orinoco basin were found to prey on larger fishes as Astyanax spp. , Brycon whitei , and Prochilodus mariae ( Rodríguez-Olarte & Taphorn, 2006) . Shrimps were also recorded in its diet ( Lasso, 2004: 157). The species presents some importance in fisheries at the río Putumayo, río Caquetá, and río Guaviare, Colombia ( Salinas-Coy & Agudelo-Córdoba, 2000; Diaz-Sarmiento & Alvarez-Léon, 2003). At the río Orinoco basin in Venezuela, the species is mostly targeted by subsistence fishermen or by sport fishermen, being uncommon in fish markets and having a low market value ( Rodríguez-Olarte & Taphorn, 2006). The species is reported to have declined considerably in some rivers of the Orinoco Andean piedmont in Venezuela ( Winemiller et al., 1996).

Remarks. The first reference in the literature to the species based on a preserved specimen was Pellegrin (1899), who identified a specimen collected at the río Apure, río Orinoco basin, Venezuela, as S. hilarii , an identification followed by several subsequent authors who identified Salminus specimens from the Orinoco and upper Amazon basin (e.g., Pearson, 1924; La Monte, 1935; Fowler, 1943; Mago-Leccia, 1970; Saul, 1975; Stewart et al., 1987; Ferreira et al., 1988; Silvano et al., 2001). Less commonly, S. iquitensis was mistakenly identified as S. affinis , an error first made by Eigenmann (1922) and followed by some subsequent authors (e.g., Eigenmann & Allen, 1942; Ortega, 1996; Goulding et al., 2003). There are even some misidentifications of S. iquitensis with S. maxillosus (a synonym of S. brasiliensis ) in the literature ( Fowler, 1945; Barriga, 1991). The confusion surrounding the identity of the Salminus species occurring at the río Orinoco and western Amazon basins can be appraised from Géry & Lauzanne (1990: 122–123), who considered S. hilarii as ocurring in the “... Purus, Haut Madeira, Haute Amazonie, Apuré, Caroni, Orteguaza” and S. affinis in the “...Haut Ucayali et Rio Mamoré”, and from Barriga (1991: 27), DiazSarmiento & Alvarez-León (2003: 313), Ortega et al. (2011), Barriga (2012: 108), and Meza-Vargas et al. (2021), who listed two to four Salminus ( S. affinis , S. hilarii , S. maxillosus , and S. brevidens ) as occurring in the eastern (Amazonian) slope of Ecuador, in the río Caquetá in Colombia, and in the Amazon basin in Peru, when all these names actually refer to a single species, S. iquitensis . The first author to notice the inadequacy of the identification of the Salminus species occurring at northern cisandean South America was Pearson (1924: 49), who stated that “The species of this genus are not yet definitely delimitated. It seems that there have been many mistaken identifications in previous records, and the identification of the present specimens may be mistaken”. A note appended to the specimen MNHN 1898-22, by an ichthyologist who studied this specimen (perharps J. Géry), noticed that “cet individu parait plus proche de affinis ( Ecuador & Cauca) que de hilarii (Rio S. Francisco)”. The historical confusion among Salminus iquitensis , S. affinis and S. hilarii can be explained by the similar, overlapping scale counts among these species, e.g., 65–87, modally 75, lateral-line scales in S. iquitensis , vs. 64–81, modally 71 in S. affinis , and 54–72, modally 65, in S. hilarii .

We herein employ the name Holobrycon iquitensis Nakashima (1941) , under the new combination Salminus iquitensis , for the species of Salminus from upper Amazon, rio Branco, and río Orinoco basins. The description and drawing by Nakashima (1941) clearly depict a Salminus and it is not clear why he chooses instead to describe it under the genus Holobrycon (currently a synonym of Brycon ; Lima, 2017). Holobrycon iquitensis was almost immediately considered as a synonym of S. hilarii (Myers in Eigenmann & Allen, 1942: 260, footnote), and was typically considered as such in the literature (e.g., Géry & Lauzanne, 1990: 116; Lima et al., 2003: 156), with the exception of Fowler (1950: 305), who considered it as a synonym of Salminus maxillosus (presently a synonym of S. brasiliensis ). However, the scale counts provided by Nakashima (1941) —72 lateral-line scales, 10 horizontal scale series between dorsal-fin origin and lateral line, and 5 horizontal scale series between lateral line and pelvic-fin insertion—as well as the origin of the specimen studied by him (“cercanias del puerto de Iquitos”) unambiguously assign the name as available for the species occurring in the upper Amazon basin, as well as the rio Branco and the río Orinoco basin. The type specimen or specimens of the species described by Nakashima (1941), including Holobrycon iquitensis , no longer exist (H. Ortega pers. comm.). We herein designate the specimen ANSP 178157, collected at the río Napo, a little downstream from Iquitos, as the neotype of Holobrycon iquitensis . The combination Salminus iquitensis has already appeared in the literature ( Buckup et al., 2007; Lehmann et al., 2009; Ortega et al., 2011; Lima, 2017; Dagosta & de Pinna, 2019), even though no formal recombination of the name Holobrycon iquitensis and revalidation of the species was available until now.

Material examined. Type material. ANSP 178157 (1, 148.2 mm SL), Peru, Loreto, Mazan, río Napo, upstream mouth of río Mazan , near Mazan , 3º29′10′′S, 73º6′24′′W; M.H. Sabaj, M.W. Littmann & K.A. Elkin, 5 Aug 2001. Neotype of Holobrycon iquitensis Nakashima , by present designation.

Not types. Bolivia: CAS 17229 (1, 301.0 mm SL), La Paz, Ixiamas (río Beni basin), c. 13º48′S, 68º6′W; N. Pearson, Dec 1921. MNHN 1989-1437 (1, 146.5 mm SL), Beni, L. Santo Antonio, mouth of río Securé, above Trinidad ( río Mamoré ), c. 15º20′S, 65º1′W; L. Lauzanne & G. Loubens, 9 June 1986. Brazil, Acre: MZUSP 50456 (1, 211.0 mm SL), Marechal Thaumaturgo, rio Juruá, Natal , c. 9º18′S, 72º41′W; R.A.S. Silvano, 12 Jul 1994. MZUSP 101091 (1, 332.7 mm SL), Marechal Thaumaturgo, rio Breu (trib. rio Juruá ), c. 9º24′S, 72º42′W; B.F. Amaral, 1995–1996. ZUEC 7454 (1, 173.2 mm SL), Cruzeiro do Sul, rio Moa, Lago do Mutuca , c. 7º37′S, 72º47′W; G. V. Andrade et al., 4 June 1982. MCZ 33530 (1, 164.4 mm SL), mouth of rio Macau ã (trib. rio Iaco, trib. rio Purus), c. 9º12′S, 68º44′W; B. Krukoff, 1933. USNM 94651 (1, 155.0 mm SL), mouth of rio Macauã (trib. rio Iaco, rio Purus basin), c. 9º12′S, 68º44′W; B.A. Krukoff, 26 Dec 1934. MCP 29029 (3, 230.1– 240.9 mm SL), Sena Madureira, rio Purus (fish market), 9°9′S, 68°35′W; L. R. Malabarba, 8 Aug 2001. Rondônia: MCP 37564 (1, 165.8 mm SL), Porto Velho, Rio Abunã, at its mouth at rio Madeira, ferryboat at road BR-364, 9°40′27′S, 65°26′46′W; P.A. Buckup, F.C. T. Lima et al., 25 July 2004. UFRO 5333 (1, 127.5 mm SL), Porto Velho, igarapé Jatuarana (trib. rio Madeira), 8º46′24′′S, 64º2′47′′W; L. Nogueira, 14 Apr 2012. UFRO 9866 (1, 194.4 mm SL), Porto Velho, igarapé Jatuarana (trib. rio Madeira), 8º50′20′′S, 64º2′44′′W; C.H. Zawadzki, 3 Jul 2009. Roraima: ZUEC 17546 (2, 224.1– 267.1 mm SL): Amajari, rio Acari (trib. rio Parimé, rio Uraricoera basin), 4º4′46′′N, 61º17′17′′W; I. Mesquita, 29 Jan 2022. INPA 10791 (1, 163.1 mm SL): Mucajaí, rio Mucajaí, mouth of igarapé Traíra, above Cachoeira Pared ã o, 2º56′23′′N, 61º37′14′′W; E.G. Ferreira & G.M. Santos, 1 Oct 1986. NMW 56870 (1, 166.6 mm SL), Caracaraí, Cachoeira do Bem Querer, rio Branco , c. 1º55′N, 61º1′W; J.D. Haseman, Oct 1912. Peru, Depto. Amazonas: CAS 54358 (4, 16.4–37.6 mm SL), río Santiago, Pinglo , c. 4º25′S, 77º39′W; D.J. Stewart, 8 April 1980. Depto. Cuzco: FMNH 76642 (1, 205 mm SL), Marcapata, Río Nusiniscato, Huajyumbe , c. 13º15′S, 70º34′W; C. Kalinowski, June 1950. Depto. Loreto: CAS 24794 (1, 138.2 mm SL), río Ucayali at Bretaña , c. 5º13′S, 74º18′W; W. R. Allen , 2–9 Sept 1920. CAS 158705 (1, 149.2 mm SL), Alto Marañon below Paztaza , c. 4º49′S, 76º19′W; W. R. Allen , Oct 1920. MUSM 10895 (1, 65.2 mm SL), río Chambira, trib. río Pintoyacu, trib. río Nanay , San Juan village , c. 4º36′S, 74º52′W; E. Castro, 14 March 1997. MUSM 18251 (3, 19.8–24.1 mm SL), Parinari, mouth of río Chambira, río Marañon basin, c. 4º36′S, 74º52′W; H. Ortega & E. Castro, 8 March 2001. BMNH 1977.3.10.307-308 (2, 129.8–139.0 mm SL), Burro Caño, near Iquitos; M. Chapman, 30 May 1974. INHS 54972 (1, 142.3 mm SL), Río Napo, opposite Mazán, N channel Río Napo , N of Isla Milagro, 3°22′59”S, 73°5′12′W; M.H. Sabaj et al., 30–31 July 1999. ANSP 178434 (1, 129.3 mm SL), Iquitos, río Nanay at Pampa Chica , 3º45′9′′S, 73º17′00′′W; M.H. Sabaj et al., 2 Aug 2001. CAS ( SU) 36907 (1, 187.0 mm SL); CAS ( SU) 68468 (1, 136.5 mm SL), Caño de Shansho, near Pebas , c. 3º21′S, 71º47′W; W.G. Scherer, Sept-Oct 1936. Depto. Madre de Dios: MUSM 7569 (4, 149.2– 187.8 mm SL), Tambopata, río Madre de Dios, near Boca Valencia , c. 12º30′S, 68º49′W; F. Chang et al., 27 Jul 1995. MUSM 16539 (1, 138.0 mm SL), Tambopata , río Madre de Dios, Playa Alta; C. Cañas, 16 Sept 1997. MUSM 8160 (1, 138.8 mm SL), río Tambopata, Santuario Nacional Pampas de Heath, río Heath , c. 13º19′S, 68º52′W; H. Ortega, 6 June 1992. MUSM 16345 (1, 160.6 mm SL), Tambopata , río Madre de Dios, Tres Islas; C. Cañas, 16 Sept 1997. MUSM 19149 (1, 157.4 mm SL), Tambopata, río Los Amigos, beach 100 m downstream Puerto CRA, c. 12º33′S, 70º5′W; M. Goulding, R. Barthem & C. Cañas, 23 Aug 2001. MUSM 20467 (1, 141.5 mm SL), Tahuamanu, río Tahuamanu, road Puerto Maldonado / Iñaparí , 11º27′47′′S, 69º18′24′′W; H. Ortega et al., 9 Aug 2003. MUSM 8283 (2, 127.0– 142.7 mm SL), Tambopata , río Madre de Dios, Gamitana; F. Chang, 15 Sept 1995. MUSM 16562 (1, 130.6 mm SL), Tambopata , rio Madre de Dios, San Francisco; C. Cañas, 15 Sept 1997. MUSM 20125 (1, 94.3 mm SL), Manu , río Madre de Dios, sand beach in front of CRA, c. 12º35′S, 70º4′W; M. Goulding et al., 3 May 2002. MUSM 20115 (1, 74.5 mm SL), Manu , río Madre de Dios, sand beach in front of CRA, c. 12º35′S, 70º4′W; M. Goulding et al., 30 April 2002. USNM 264008 (1, 250.0 mm SL), stream trib. río Tambopata, upstream mouth of río La Torre , 12º49′40′′S, 69º18′W; R. P. Vari, H. Ortega & S.L. Jewett, 23 Aug 1983. USNM 263995 (1, 133.5 mm SL), río Tambopata, upstream 200 m above río La Torre , c. 12º50′S, 69º18′W; R. P. Vari & H. Ortega, 21 Aug 1983. USNM 264007 (2, 148.7– 179.6 mm SL), Río Madre de Dios, 10 km downstream junction with río Tambopata , c. 12º36′S, 69º4′W; R. P. Vari, H. Ortega & S.L. Jewett, 25 Aug 1983. ROM 64026 (1, 76.6 mm SL); USNM 319363 (1, 78.4 mm SL), Manu, Manu National Park, Pakitza , quebrada Pica Flor , 11º56′40′′S, 71º16′59′′W; H. Ortega, 21 Apr 1991. ANSP 180556 (1, 119.1 mm SL), río Madre de Dios, Isla Rolin, downstream Puerto Maldonado, 12º33′50′′S, 69º3′1′′W; M.H. Sabaj, N. Salcedo, L. Sousa, M. Arce et al., 29 Jul 2004.

Depto. Pazco: CAS 79293 (1, 382.2 mm SL), río Azupirazu (trib. río Palcazu, trib. río Pachítea ) at Tambo , c. 10º20′S, 74º54′W; W. R. Allen , 8–9 Jul 1920 . MUSM 33637 (1, 242.0 mm SL), Oxapampa, Izcozocín, Yanachaga- Chemillén National Park , río Izcozacín , quebrada Pescado , 10º22′38′′S, 75º14′34′′W; M.H. Hidalgo & M. Rojas, 16 Sept 2007 . USNM 229054 (1, 266.0 mm SL), río Palcazu (trib. río Pachitea), at Iscozacin , 10º11′S, 75º9′W; P. Bay- ley, 25 Jul 1981 . Depto. Ucayali : MZUSP 26078 (1, 250.0 mm SL), río Huacamayo, road Aguaytia-Pucallpa , km 7.5, c. 9º0′S, 75º29′W; J. Guevara, 11 Oct 1971 . MZUSP 25949 (4, 1 c&s, 122.8–142.2 mm SL), río Ucayali, Masisea, prov. Cel. Portillo , c. 8º35′S, 74º20′W; H. Ortega, 15 Aug 1975 . MUSM 29 (2, 235.8– 250.6 mm SL), río Huacamayo, trib. río Aguaytia, km 150, road Pucallpa / Huánuco , c. 9º0′S, 75º29′W; J. Guevara, 14 Oct 1971 . MUSM 30 (5, 124.6– 195.2 mm SL), Pucallpa, río Ucayali , San Juan; H. Ortega, 26 May 1983 . MUSM 12579 (1, 162.4 mm SL), Coronel Portillo, río Ucayali, Boca del Pau , near Pucallpa , c. 8°23′S, 74°33′W; H.W. Koepcke, 11 June 1958 . MUSM 109 (1, 136.5 mm SL), Pucallpa, Masisea, Lobococha , c. 8º35′S, 74º20′W; H. Ortega, 20 April 1983 . MUSM 4240 (3, 130.2– 149.1 mm SL), Pucallpa, río Ucayali, Utiquínia , c. 8º13′S, 74º20′W; H. Ortega, 14 June 1981 . USNM 261514 (1, 134.6 mm SL), río Ucayali , Nuevo San Juan, Pucallpa; H. Ortega, 26 May 1983 . MUSM 56 (2, 144.9– 161.4 mm SL), río Chio, road Pucallpa / Tingo Maria; H. Ortega & P. de Rham, 6 Nov 1984 . MUSM 730 (1, 141.2 mm SL), Pucallpa, río Ucayali , c. 8º24′S, 74º31′W; H. Ortega, 22 Aug 1984 . MUSM 577 (1, 122.2 mm SL), Pucallpa, río Neshuya , c. 8º38′S, 74º58′W; G. Contreras & C. Rojas, 24 July 1981 . MUSM 5535 (1, 134.5 mm SL), Pucallpa, río Ucayali , Shanahao; H. Ortega, 7 May 1975 . MUSM 7445 (1, 100.0 mm SL), Yarina Yarinococha, Caño de Panaillo ; C. Riofrio et al., 16 June 1995 . CAS 24793 (2, 202.0–215.0 mm SL), río Ucayali, sandbars at Orellana, below Contamana , c. 6º55′S, 75º9′W; W . R. Allen , 22–23 Aug 1920 . Ecuador: FMNH 117245 (1, 315.0 mm SL), Napo, río Coca, below río Sardinas mouth, c. 0º6′S, 77º12′W; D.J. Stewart et al., 28 Sept 1981 . ANSP 130599 (1, 204.0 mm SL), Napo, río Conejo, Santa Cecilia , c. 0º6′N, 76º51′W; W.J. Saul, 10 June 1968 . Colombia, río Amazonas basin, Depto. Putumayo: ICNMHN 14482 (2, 149.8–151.0 mm SL), Puerto Leguízamo , Puntales , c. 0º11′S, 74º48′W; M. Leiva, 10 Nov 2005 . CIACOL 5355 (1, 157.0 mm SL): Puerto Leguízamo, La Tagua, stream trib. río Caquetá, 0º5′6′′S, 74º40′39′′W; A.A. Santos, 9 March 2021 . MLS 103 (1, 204.0 mm SL), Puerto Asis, río Putumayo basin, c. 0º30′N, 76º31′W; Nicéforo Maria, Feb 1938 . Depto. Caquetá: IAvH-P 5801 (1, 228.0 mm SL), río Caquetá, Araracuara , c. 0º37′S, 72º23′W; G. Gutierrez, 16 Feb 1981 . ANSP 95869 (1, 235.0 mm SL), Florencia, río Orteguasa basin, c. 1º36′N, 75º37′W; Brother Nicéforo Maria, 1932. CIACOL 1532 (1, 285 mm SL), Florencia, quebrada La Carbona, km 15 road Florencia-Neiva , 1º42′25′′N, 75º37′22′′W; J.D. Bogotá-Gregory, 28 Jul 2013 . MLS 104 (1, 178.3 mm SL), Milán, río Orteguaza, Villa Fátima (= Caserío La Rastra ); Nicéforo Maria, 27 Aug 1951 . ANSP 169622 (1, 245 mm SL), Morelia (río Caquetá drainage), c. 1º29′N, 75º44′W; K. Von Sneidern, 1945. CIACOL 1796 (1, 298 mm SL): San José del Fragua, río Fragua Grande (trib. río Caquetá), 1º8′38′′N, 76º16′58′′W; L. Penha, 10 Jun 2015 . Depto. Amazonas: IAvH-P 1801 (1, 175.0 mm SL), La Chorrera, Quebrada La Tag ̧a, río Igara-Paraná , c. 1°9′S, 72°33′W; C.A. Santamaría, 29 Jul 1994 . Río Orinoco basin: Depto. Casanare: IAvH-P 930 (2, 140.5– 144.2 mm SL), río Túa , ca. 10 km above mouth into río Meta, c. 4°25′N, 72°33′W; R. Scully & F. Garzón, 28 Nov 1978 . IAvH-P 1940 (2, 230.9–260.0 mm SL), Aguazul, Caño Grande , c. 4°51′N, 72°54′W; J. Jiménez, 8 Jul 1992 . IAvH-P 2906 (1, 245.0 mm SL), Tauramena, río Cusiana , c. 4°31′N, 71°51′W; F. Rodríguez, 11 March 1993 . IAvH-P 3676 (1, 176.2 mm SL), Tauramena, río Caja, trib. río Cusiana , c. 5°0′N, 72°42′W; V. Ortíz , 14 Nov 1994 . IAvH-P 3677 (7, 124.4– 158.9 mm SL), Tauramena, río Túa , c. 4°18′N, 72°36′W; F. Rodríguez, 1 Aug 1993 . IAvHP 3678 (1, 138.4 mm SL), Tauramena, río Túa , c. 4°18′N, 72°36′W; V. Ortíz , 13 Feb 1996 . IAvH-P 3681 (6, 156.7– 206.0 mm SL), Tauramena, río Cusiana and río Túa , c. 4°31′N, 71°51′W; V. Ortíz , 1 May 1995 . IAvH-P 3333 (1, 212.8 mm SL), Pore, río Pauto , c. 5°9′N, 70°54′W; L. Narváez, 25 Feb 1993 . IAvH-P 3680 (2, 199.0–237.0 mm SL), Pore, río Pauto , c. 5°9′N, 70°54′W; V. Ortíz , 1 Feb 1996 . MPUJ 8820 (1, 217.8 mm SL), Pore, Brisas del Pauto, río Pauto , 5º32′59′′S, 71º54′57′′W; J. Zamudio, 18 March 2015 . IAvH-P 3675 (1, 187.8 mm SL), Yopal, río Charte , c. 4°42′N, 72°10′W; no collector specified, March 1993 . IAvH-P 3679 (4, 163.0–223.0 mm SL), Yopal, río Cravo Sur, río Payero and río Tocaria , c. 4°42′N, 71°36′W ; V. Ortíz , 7 March 1994 . ICNMHN 13302 (1, 174.6 mm SL), Maní, río Cuarte, 9 km from Maní (río Meta basin), c. 4º50′N, 72º14′W; P. Cala, 19 Feb 1971 . MPUJ 17029 (1, 215.3 mm SL), Sabana Larga, río Upía, El Secreto bridge, 4º49′13′′N, 73º4′23′′W; T.P. Carvalho et al., 22 Feb 2022 . MPUJ 9388 (1, 235.7 mm SL), Tamara, Caño Mochila (trib. río Pauto ), near Aguas Termales , 5º44′21′′N, 72º12′30′′W; J. Zamudio, 20 Aug 2015 . MPUJ 9389 (1, 295.8 mm SL), Tamara, Quebrada Bayagua (trib. Quebrada Honda ), 5º47′57′′N, 72º6′41′′W; J. Zamudio, 18 Aug 2015 . MPUJ 11868 (1, 133.1 mm SL), Paz de Ariporo, Caño La Her- mosa, 5º42′21′′N, 71º1′11′′W; A. Urbano-Bonilla, 23 Sept 2016 . IAvH-P 13622 (1, 168.0 mm SL), Hato Corozal, río Ariporo, vereda La Chapa , 5°54′10”N, 71°29′15”W; M. Roa, J.G. Albornoz-Garzón & D. Montoya, 25 Nov 2014 . CZUT-IC 12860 (3, 168.1– 185.2 mm SL), río Ariporo, Correjimiento La Chapa, Caño Aceites , 5º54′14′′N, 71º29′16′′W; D. Montoya, 25 Nov 2014 . MPUJ 16809 (1, 166.8 mm SL), Hato Corozal, río Chire (trib. río Casanare), 6º12′17′′N, 71º50′31′′W; T.P. Carvalho et al., 16 Feb 2022 . MPUJ 16673 (1, 220.0 mm SL), Hato Corozal, río Casanare, Fínca San Joaquin , 6º12′19′′N, 71º28′38′′W; T.P. Carvalho et al., 14 Feb 2022 . IAvH-P 3682 (3, 196.4– 273.0 mm SL), no specific locality; V. Ortíz, 18 Nov 1995 . Depto. Cundinamarca: MLS 1619 (1, 135.1 mm SL), Cundinamarca, Medina, stream near Medina between Cumaral-Medina , c. 4º29′N, 73º20′W; E.C. Bonilla, 10 Oct 1998 . Depto. Guaviare: CZUT-IC 7147 (2, 133.6– 205.3 mm SL), San José del Guaviare, Caño El Flauta (trib. río Guayabero ), 2º20′19′′N, 73º2′38′′W; A. Urbano-Bonilla, 9 Sept 2010 . Depto. Meta: IAvH-P 1049 (1, 243.0 mm SL), Puerto López, río Humea , near mouth of río Negrito , c. 4º9′N, 72º54′W; R. Scully , 10 Dec 1979 . IAvH-P 2569 (1, 149.5 mm SL), Puerto López, río Meta, near the mouth of río Yucao , c. 4°27′N, 72°6′W; M.C. Blanco, no date. ICNMHN 1730 (3, 140.8– 165.7 mm SL), Puerto López, río Metica , laguna de Menequa, c. 4º6′N, 72º56′W; G. Galvis, 8 Aug 1982 . ICNMHN 1281 (1, 116.1 mm SL), Puerto López, Laguna de Menegua , c. 4º6′N, 72º56′W; G. Galvis et al., 11 Aug 1985 . MPUJ 2943 (1, 215.2 mm SL), San Martín, Vereda Montebello, río Caño Camoa , 3º39′48′′N, 73º36′34′′W; S. Prada-Pedreros et al., 9 Apr 2007 . IAvH-P 1185 (1, 104.9 mm SL), Río Guayabero , playa La Danta, c. 2°36′N, 72°46′W; M.C. Blanco, 30 Oct 1983 . CAS ( SU) 53760 (1, 159.3 mm SL), Cordillera Macarena, small brook into río Guayabero , 5 mi. below El Refugio, c. 2º17′N, 73º44′W; T.D. White & G.S. Myers, 24 Feb 1960 . USNM 181364 (3, 149.0–195.0 mm SL), río Guaviare, “ Finca of Captain Series ”; S. Weinstein, Dec 1957 . FMNH 73460 (2, 198.0– 205 mm SL), río Guaviare (no precise locality); S. Weinstein, Dec 1957 . Depto. Vichada: CZUT-IC 13540 (1, 216.3 mm SL), río Cadá, Vereda los Mangos , 3º37′30′′N N, 69º51′35′ W; E. Sanchez, 14 Jan 2015 . ICNMHN 3889 (12, 58.3–76.3 mm SL), Puerto Carreño, río Meta (rebalse), c. 6º12′N, 67º28′W; J. Maldonado et al., 23 Jun 1998 . Venezuela, río Orinoco basin: Estado Amazonas: CAS ( SU) 24795 (2, 157.5– 176.3 mm SL), río Orinoco bifurcation, Tamatama , c. 3º9′N, 65º51′W; C. Ternetz, 20 March 1925 . ANSP 160015 (1, 157.6 mm SL), río Meta, 20 min. upstream confluence with río Orinoco , c. 6º15′N, 67º37′W; B. Chernoff et al., 27 Nov 1985 . ANSP 160011 (1, 197.8 mm SL), Caño Orera at junction of río Orinoco, ca. 15 min. upstream from El Burro , c. 6º11′N, 67º26′W; A. Machado-Allison et al., 27 Nov 1985 . MZUSP 54584 (2, 140.5– 151.9 mm SL), río Orinoco, opposite to El Burro , c. 6º12′N, 67º26′W; A. Machado, C. Marrero & L. Aguana, 28 Nov 1985 . ANSP 160012 (3, 130.4–170.0 mm SL) ; ANSP 160013 (1, 179.2 mm SL), río Orinoco at El Burro, c. 6º12′N, 67º26′W; B. Chernoff et al., 26 Nov 1985 . ANSP 160014 (6, 127.0– 187.9 mm SL), río Orinoco at El Burro, c. 6º21′N, 67º26′W; L. Aguana, 26 Nov 1985 . Estado Apure: MCNG 43810 (2, 130.8– 186.6 mm SL), Hato El Frío , c. 7º50′N, 68º54′W; C. Lasso et al., 15 Nov 1990 . MNHN 1898-22 (1, 105.8 mm SL), Apure (no precise locality); M.F. Geay, 1893– 1895. Estado Aragua: USNM 121399 (1, 173.0 mm SL), río Guarico and tribs. between San Sebastian and San Casemiro , c. 9º56′N, 67º3′W; L.P. Schultz, G. Zuloage, W. Phelps & R. Sherman, 14 Apr 1943 . Estado Barinas: USNM 194213 (3, 195.0–264.0 mm SL), río Nitiado-Seco (not located); P. Bottome & W. Wallis, 14 Sept 1964 . USNM 194208 (2, 202.0–240.0 mm SL), río Las Palmas (trib. río Paguey ), c. 8º4′N, 70º20′W; P. Bottome & W. Wallis, 15 Jul 1958 . USNM 311073 (1, 245.0 mm SL), río Corozo, on Barinas/ Pedraza road, between río “Paguzy” (= Paguey ?) and Barinas, c. 8º34′N, 70º22′W; P. Bottome & W. Wallis, 16 Jul 1958 . Estado Bolivar: ANSP 160010 (1, 142.9 mm SL), caño 15.1 km E of río Parguaza, ferry crossing on Caicara-Puerto Ayacucho hwy, 6º26′28′′N, 67º9′24′′W; B. Chernoff et al., 28 Nov 1985 . Estado Guarico: FMNH 85447 (1, 188.0 mm SL), río Manapire , 113.1 km S of Chaguaramas 2.5 km E of highway, c. 8º20′N, 65º7′W; J. Thomerson, D. Hicks, D.C. Taphorn & D. Greenfield, 5 Jan 1975 . UMMZ 214816 (1, 149.2 mm SL), Guarico dam near Calabozo , río Guarico drainage, c. 8º56′30′′N, 67º24′30′′W; W. Fink et al., 22 Feb 1987 . Estado Lara: MCNG 34962 (1, 171.2 mm SL), Parque Nacional Yacambu, río Guaical , c. 9º41′N, 69º32′W; C. Marrero et al., 16 March 1994 . Estado Portuguesa: MCNG 35450 (1, 224.1 mm SL), Caño Volcán, road between Tucupido and dam, 8º59′15′′N, 69º53′30′′W; K. Winemiller et al., 7 June 1997 . MCNG 1723 (1, 190.3 mm SL), Caño El Diablo, above Guanarito, at its mouth at río Guanare , c. 8º43′N, 69º21′W; S. Reid & D. Taphorn, 10 Sept 1980 . MCNG 3514 (1, 209.3 mm SL), río Portuguesa at El Ríncon, 14 km east from Papelon , c. 8º54′N, 69º22′W; D. Taphorn, 28 Feb 1979 . INHS 59899 (1, 132.6 mm SL), río Are (trib. río Guache, río Portuguesa drainage), Aparicion , at hwy. 5 bridge, 9º22′54′′N, 69º23′8′′W; L.M. Page et al., 23 Dec 1990 .

Incorrect locality: USNM 100771 (1, 175.0 mm SL), “ Colombia, Norte de Santander, Río Pamplonita, near Cucuta; Brother Niceforo Maria, 1935” (see Distribution , above) .

Figures

FIGURE 12. Salminus iquitensis: (A) ANSP 178157, 148.2 mm SL, Peru, Loreto, río Napo; neotype of Holobrycon iquitensis Nakashima (photo by M.H. Sabaj); (B) MZUSP 26078, 250 mm SL, Peru, Ucayali, río Huacamayo; (C) MCNG 3514, 209.3 mm SL, Venezuela, Portuguesa, río Portuguesa; (D) IAvH-P 3680, 237.0 mm SL; Colombia, Casanare, río Pauto.

FIGURE 13. Salminus iquitensis, living specimens: (A) Peru, Ucayali, río Aguaytia; (B) ROM 93073, Ecuador, Paztaza, río Bobonaza; (C) CZUT-IC 7147, 133.6 mm SL: Colombia, Guaviare, Río Guayabero basin; (D) IAvH-P 17096, Colombia, Meta, Río Guayabero. Photo A by A. Linares, B by N.K. Lujan, C by A. Urbano-Bonilla, D by Mark Sabaj.

FIGURE 14. Map of northern South America, showing the distribution of Salminus iquitensis (red dots; red star indicates the type-locality; purple dot indicates a locality at the rio Branco basin from which no voucher specimen is available; see item “Distribution” of S. iquitensis).