Brycon amazonicus ( Agassiz, 1829 ),

Lima, Flávio C. T., 2017, A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae), Zootaxa 4222 (1), pp. 1-189: 112-129

publication ID 10.5281/zenodo.257769

publication LSID

persistent identifier

treatment provided by


scientific name

Brycon amazonicus ( Agassiz, 1829 )


Brycon amazonicus ( Agassiz, 1829) 

( Figs. 66–73View FIGURE 66View FIGURE 67View FIGURE 68View FIGURE 69View FIGURE 70View FIGURE 71View FIGURE 72View FIGURE 73)

Chalceus amazonicus Agassiz, in Spix & Agassiz, 1829: 68  –69, pl. 35 (Type locality: “in fluvio Amazonum”); Howes, 1982: 9 (comments).

Chalceus carpophaga Valenciennes, in Cuvier & Valenciennes, 1850: 252  –253 (Type locality: “l’ Esséquibo ” “l ´ Amazone ”, retricted by Géry & Mahnert (1992: 816) to “ Essequibo ”). [not Kner, 1860: 12].

Chalceus hilarii  (not Valenciennes): Kner, 1860: 10 –11 (part; “ Salto Theotonio”).

Megalobrycon cephalus Günther, 1869: 423  –424, fig. 1 (Type locality: “Upper Amazons”); Regan, 1905: 190 (Rio Negro; based on Wallace’s drawing).

Brycon capito Cope, 1872: 261  –262 (Type locality: “Ambyiacu”); Fowler, 1906: 446, fig. 42 (holotype, redescription).

Brycon longiceps Steindachner, 1879a: 150  (Type locality: “Orinoco bei Ciudad Bolívar”); Steindachner, 1879b: 157 –158, pl.1, fig.5 (full description); Mago-Leccia, 1970: 69 (listed, Venezuela; common name); Lasso, 1988: 127 (Lower Río Orinoco, Laguna los Barrancos; common name); Lima, 2003: 175 (primary type material; as a synonym of B. amazonicus  ).

Brycon siebenthalae Eigenmann, 1912: 372  , pl. 54, fig. 3 (Type locality: “Mud Creek, Aruka River”); Géry, 1977: 338 (premaxillary teeth arrangement, holotype); Howes, 1982: 44 (comments); Cala, 1986: 91 (Río Ariari, Río Tomo; Río Guaviare; Río Vichada; western llanos of Colombia; common name; economic importance); Useche-L. et al., 1993: 341 – 348, 350–351 (Río Cafre, trib. Río Guaybero, Depto. Meta, Colombia; breeding; length/weight relationship; condition factor; gonadossomatic index; fecundity; feeding habits); Bernal-Ramírez & Cala, 1997: 55 –63 (Río Guayabero, Río Guaviare basin, Colombia: diet, seasonality); Lima, 2003: 175 (primary type material; as a synonym of B. amazonicus  ).

Brycon pellegrini Holly, 1929: 208  –209 (Type locality: “Manaos”); Howes, 1982: 41 (possible synonym of Brycon cephalus  ); Lima, 2003: 175 (primary type material; as a synonym of B. amazonicus  ).

Chalceus taeniatus  (not Jardine, in Jardine & Schomburgk): Magalhães, 1931: 130 (Amazonas; common name, natural history).

Brycon siebenthalae iquitensis Nakashima, 1941: 69  –70, fig. (Type locality: “cercanías del puerto de Iquitos”).

Brycon hilarii  (not Valenciennes): Amaral-Campos, 1950: 140 (in part; “ Pará, rio Amazonas ”); Marlier, 1968: 56 (Paraná da Eva, central Amazon).

? Brycon stubelli  (not Steindachner): Amaral Campos, 1950: 142 (rio Juruá, Amazonas).

Brycon  sp.: Smith, 1979: 131 (Amazonas, Itacoatiara); Goulding, 1979: 95 –100, 133, 145–149 (Rio Machado, Rondônia; migrations, fisheries); Goulding, 1980: 68 –73 (Rio Machado, Rondônia; common name, diet, migrations); Crampton, 1999: 15, 27 (Reserva Mamirauá, lower rio Japurá, Amazonas); Barthem, 1999: 82, 94 (Central Amazon; fishery); Toledo- Piza, 2002: 122 –123 (drawing by A.R. Wallace; Yavita, rio Orinoco, Venezuela); Fernández et al., 2006: 58 (photo; upper Río Cataniapo, Rio Orinoco basin, estado Amazonas, Venezuela).

Brycon carpophagus: Howes, 1982: 15  –18 (in part; syntypes); Géry & Mahnert, 1992: 815 –817 (lectotype designation; Brycon siebenthalae  as a synonym); Lima, 2003: 175 (primary type material; as a synonym of B. amazonicus  ). [Not Mérona et al., 2001: 387, 391].

Brycon cephalus: Howes, 1982: 18  –20 (in part; Brazil, Manaus; Monte Alegre; “Manacapuni”); Zaniboni Filho et al., 1988: 41–50, fig. 7 (Brazil, rio Negro, rio Solimões, rio Japurá and rio Juruá; discussion); Zaniboni Filho & Resende, 1988: 833– 844 (gonad anatomy, maturity scale, type of spawning); Géry & Mahnert, 1992: 794 –800, 802, figs. 1 (lower), 5 (Central Amazon; diagnosis; common name; comparison with B. melanopterus  ); Ferreira et al., 1998: 93, fig. 61 (Pará, rio Tapajós, Santarém; common name; fisheries); Pizango-Paima et al., 2001: 509 –520 (diet, seasonality, Central Amazon); Araújo- Lima & Ruffino, 2003: 253 –254 (migrations, fisheries; Amazon basin, Brazil); Diaz-Sarmiento & Alvarez-Léon, 2003: 311 (Amazon basin, Colombia; fisheries); Petry et al., 2003: 565, 575 (Ilha de Marchantaria, rio Solimões; occurrence related to macrophytes and other environmental gradients); Mérona & Rankin-de-Mérona, 2004: 77 (Lago do Rei, Ilha do Careiro, rio Amazonas; diet); Lima & Araújo-Lima, 2003: 788 –790, 792–793, 795, 797 (distribution of larvae and juveniles, central Amazon); Galvis et al., 2006: 188 –189 (Colombia, Rio Amazonas at Leticia; short description, drawing). [Not Galvis et al., 2006: 457, photo].

Brycon cf. melanopterum  (not Cope): Braum, 1983a: 355 –371, fig. 1a (lip extension during hypoxia).

Brycon cf. melanopterus  (not Cope): Braum, 1983b: 268 –269 (lip extension during hypoxia); Werder, 1983: 445 –461 (age estimates through circuli analysis); Junk et al., 1983: 408, 414, 426–427 (Lago Camaleão, Ilha da Marchantaria, rio Solimões; abundance, morphological adaptations to endure hypoxia); Werder & Soares, 1984: 398 –416 (Manaus; age determination thorugh circuli; scale variation); Piedade et al., 2006: 1176 (ingestion of Astrocaryum jauari  fruits in the lower Rio Negro, Brazil).

Brycon melanopterus  (not Cope): Bayley, 1988: 131 –133, 136 (in part; rio Solimões floodplains, Manaus area; growth rates of young specimens related to density and seasonality); Lasso, 1992: 11, 22, fig. 2 (río Suapure, Serranía de Los Pijiguaos, Rio Orinoco basin, Est. Bolivar, Venezuela).

Brycon cf. cephalus  (not Günther): Goulding et al., 1988: 124 (Rio Negro); Ferreira, 1993: 56, 63, 86 (Pará, rio Trombetas basin: Oriximiná; igarapé Caxipacoré; rio Cachorro; rio Mapuera); Machado-Allison et al., 1999: 65 (Rio Caura, Estado Bolivar, Venezuela); Almeida-Toledo et al., 1996: 36, 39 (karyotype).

Brycon erythropterum  (not Cope): Goulding et al., 1988: 124 (Rio Negro).

Brycon amazonicus  : Lima, 2003: 175 (new combination; synonymic list, distribution, maximum length); Goulding et al., 2003: 138 (picture; rio Madre de Dios basin); Santos et al. 2006: 39 (retouched picture; description, biology, common name, fishery importance; Manaus area); Santos Filho & Batista, 2009: 195 –203 (growth, mortality rates; rio Madeira; rio Purus; Central Amazon); Lima & Ribeiro, 2011: 151, 161(“lowland” distributional pattern; influence of blackwater and tidal movements on distribution in Amazon basin); Lima et al., 2013: 228 –229 (Brazil, Rondônia, rio Madeira basin; distribution in the rio Madeira basin, short description, photo); Phillip et al., 2013: 8, 16 (Trinidad southern coast; occurrence as a vagrant species).

Brycon amazonicum: Leite, 2004: 661  –664 (Ilha da Marchantaria, rio Solimões, Amazonas; diet of juveniles). [Not Mérona et al., 2010: 106, 194]

Brycon bicolor  (not Pellegrin): Fernández et al., 2006: 54 (photo; Río Cataniapo basin, Rio Orinoco basin, estado Amazonas, Venezuela).

Brycon cf. falcatus  (not Müller & Troschel): Fernández et al., 2006: 55 (photo; upper Río Cataniapo, Rio Orinoco basin, estado Amazonas, Venezuela).

Diagnosis. Brycon amazonicus  is distinguished from all its congeners, except B. falcatus  , B. melanopterus  , B. whitei  , B. orbygnianus  , B. orthotaenia  , B. hilarii  , and B. gouldingi  by the possession of the fifth infraorbital bone higher than wide (vs. fifth infraorbital bone about as high as wide, or wider than high in the remaining species), and the presence of several narrow, longitudinal stripes along the dorsolateral surfaces of the body (vs. no narrow, longitudinal stripes along the dorsolateral surfaces of the body). It can be distinguished from B. falcatus  and B. melanopterus  by the possession of wavy longitudinal stripes along the dorsolateral surfaces of the body (vs. straight longitudinal stripes along the dorsolateral surfaces of the body), and darkened pectoral and pelvic fins (vs. pale pectoral and pelvic fins). Brycon amazonicus  is distinguished from B. orbygnianus  , B. hilarii  , B. orthotaenia  , and B. whitei  by the lack of a broad, midlateral stripe along the caudal peduncle and middle caudal-fin rays (vs. presence of such a stripe), and darkened pectoral and pelvic fins (vs. pale pectoral and pelvic fins). The species most similar to Brycon amazonicus  is B. gouldingi  , from which it can be distinguished by the presence of dark pigment on the caudal peduncle and caudal fin diffuse, never crescent- or V-shaped (vs. a distinct crescent- or Vshaped blotch on the caudal peduncle and caudal fin). See the item “Comparisons”, below, for more details on the diagnosis between both species, as well as a discussion on the similarity in color pattern between Brycon amazonicus  specimens from the Amazon basin in Peru with specimens of the partially sympatric B. hilarii  .

Description. Morphometric data are presented in Table 18. Large-sized species, largest examined specimen 485.0 mm SL. Body moderately slender to moderately high. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, straight to slightly convex from latter point to basis of supraoccipital process, moderately convex from latter point to dorsal-fin origin, straight along dorsal-fin basis, and straight to slightly convex from dorsal-fin basis to adipose-fin origin. Dorsal profile of caudal peduncle slightly concave. Ventral profile slightly convex from lower lip to pelvic-fin insertion, straight to slightly convex from this point to anal-fin origin and approximately straight along anal-fin base. Ventral profile of caudal peduncle slightly concave.

Head profile slightly acute anteriorly, mouth terminal. Jaws approximately isognathous to slightly anisognathous, outer row of premaxillary teeth partially exposed when mouth is closed. Maxillary moderately long, extending posteriorly to anterior third of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Eight (6), 9 (16), 10 (32), 11 (33), 12 (21), 13 (7), or 14 (1) relatively small tricuspidate teeth in outer series. Three (2), 4 (28), 5 (52), 6 (30), or 7 (1) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 3 (15), 4 (84), 5 (13), or 6 (1) tricuspidate teeth between the first and third rows. Two teeth in third premaxillary row, medial teeth largest, symphyseal teeth smaller, slightly tilted towards each other, medial teeth hexa- to heptacuspidate, symphyseal teeth penta- to hexacuspidate. Maxillary margins approximately parallel, straight in profile. Twelve to 24 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 8 (11), 9 (13), 10 (16), 11 (4), 12 (5), 16 (1), or 19 (1) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, penta- to hexacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, penta- to unicuspidate. Inner (lingual) series consisting of a small, single unicuspid symphyseal tooth, situated immediately posterior to symphyseal dentary teeth of main series, plus row of 11–16 small, aciculated, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of fifth main series dentary teeth.

Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Fifty-seven (1), 61 (2), 62 (4), 63 (7), 64 (9), 65 (12), 66 (12), 67 (13), 68 (14), 69 (11), 70 (14), 71 (5), 72 (8), 73 (2), 74 (4), or 75 (2) scales in lateral line series. Laterosensory tube simple in specimens smaller than 100 mm SL, ramified in specimens larger than 100 mm SL. Tubules ramification increasing in complexity along ontogeny, specimens up to 150 mm SL with tubules with two or three branches, three to six branches in specimens between 150–250 mm SL, and with more than 10 branches and developing a dendritic pattern of ramification, with tubules often overlapping each other in larger (> 300 mm SL) specimens. Horizontal scale rows between dorsal-fin origin and lateral line 11 (12), 12 (18), 13 (56), 14 (30), or 15 (2). Horizontal scale rows between lateral line and pelvic-fin 5 (1), 6 (6), 7 (38), 8 (53), 9 (18), or 10 (1). Circumpeduncular scales 17(2), 18 (11), 19 (17), 20 (32), 21 (31), 22 (17), 23 (7), or 24 (1).

Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 11th (1), 12 th (2), or 13 th (5) vertebrae. Anal-fin rays iii (not including first, small unbranched ray only visible in cs specimens), 20 (10), 21 (17), 22 (26), 23 (24), 24 (32), 25 (9), or 26 (1). First anal-fin pterygiophore inserting behind haemal spine of 24th (1), 25th (1), or 26th (3) vertebrae. Anal-fin rays decreasing only slightly in size towards anal-fin end. Sheath of scales covering basis of anal-fin rays composed of four scale rows, lower scale row formed by 20–25 rectangular scales. Pectoral-fin rays i, 12 (5), 13 (29), 14 (71), or 15 (15). Pelvic-fin rays i, 6 (4), 7 (103), or 8 (13). Main caudal-fin rays 10/9. Caudal fin slightly forked, distal margin slightly concave. Laterosensory tube extending over interradial membrane between upper and lower caudal-fin lobes to the distal portion of fin. Laterosensory tube on caudal fin with dorsally and ventrally oriented side branches across its length.

Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First branchial arch with 13 (2), 15 (11), 16 (15), or 17 (2) lower, 1 at angle, and 13 (2), 14 (2), 15 (11), 16 (16), 17 (3), or 18 (1) upper gill rakers. Vertebrae 46 (1), 47 (1), or 48 (2). Supraneurals 8(3), 9(3), or 10 (2).

Coloration in alcohol. Top of head, snout, supraorbital, and sixth infraorbital light- to dark-brown. Dorsal portion of body light-brown to dark-brown. Second, third, fourth, and fifth infraorbitals, and opercle silvery in specimens that retained guanine, light-brown in specimens that lost this pigment due to a long storage in formalin. Dentary, maxillary, gular area, and lower portion of body light-brown. Lateral portion of body light brown, with a silvery hue. Humeral blotch present, conspicuous, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of second to third, extending longitudinally to posterior margin of sixth lateral line scales, and vertically one and half scales high. Dark, wavy longitudinal stripes formed by dark pigment concentrated on upper and lower scale margins extending along trunk moderately to highly conspicuous. Stripes more discernible dorsally. Series of irregular, narrow vertical stripes present in small (up to 80 mm SL) specimens. Diffuse dark pigmentation present at anal-fin basis, extending as a diffuse stripe into caudal-peduncle and upper caudal-fin lobe, forming an oblique stripe. Pigmentation on caudal peduncle and upper caudal-fin lobe generally less conspicuous and not forming an oblique stripe in specimens from the Río Orinoco basin, which possess instead a diffuse dark pigmentation on caudal fin. Juveniles (up to 80 mm SL) with a large, slightly assymetrical caudal peduncle extending into middle and upper caudal-fin rays basis, and anal-fin basis clear, without diffuse dark stripe. Most specimens from the upper Amazon basin in Peru (e.g., MZUSP 15266, MUSM 41154, MUSM 108, MUSM 315) with dark pigmentation concentrated on middle portion of caudal peduncle and middle caudal-fin rays. Pectoral and pelvic-fins generally dark in larger (> 150 mm SL) specimens. Lower caudal-fin rays, dorsal and anal-fins clear, with a variable amount of interradial dark pigmentantion. Adipose-fin light- to dark-grey.

Coloration in life. Based in pictures of fresh collected specimens from rio Guaporé (Rondônia), rio Sucunduri (Amazonas), lower rio Tapajós ( Ferreira et al., 1998: 93, fig. 61), rio Madre de Dios basin ( Goulding et al., 2003: 138), rio Gueppi (Loreto, Peru), and rio Solimões (Amazonas, Brazil). Overal color pattern light-grey, darker dorsally, with a silvery/plumbeous hue. Branchiostegal rays and lower half of opercle orangish in some specimens. Dark markings on anal and caudal fins and longitudinal wavy dark stripes generally very conspicuous, except in specimens collected in muddy/white waters, which generally present a drab overall coloration. All fins, but specially anal- and caudal-fins, pinkish.

Variation. There is relatively little geographical variation within Brycon amazonicus  , except that, as discussed in the item “Coloration in alcohol”, most specimens from Peruvian Amazon present dark pigmentation concentrated on the middle portion of the caudal peduncle and caudal-fin, forming a dark stripe. These specimens are attributed to Brycon amazonicus  due to their overall body shape and pectoral and pelvic fin dark coloration, and were often found in the same localities with the similar-looking B. hilarii  (see the item “Comparisons”, below).

Common names. Brazil, Manaus: “matrinchã”, “ matrinchão  ” ( Borges, 1986, Zaniboni et al., 1988; Santos et al., 2006: 39); rio Madeira basin, Rondônia: “jatuarana” ( Goulding, 1979, 1980). The same common names are applied, but with an inverse usage, to Brycon melanopterus  on those same regions (see under “Common names” of B. melanopterus  ). Venezuela: “palambra”, “bocón” ( Mago-Leccia, 1970: 69); “äi” (Piaroa language) ( Fernández et al., 2006: 54). Colombia: “yamú”, “bócon” ( Cala, 1986).

Distribution. Widespread in northern cis-andean South America, in the Rio Amazonas and Rio Orinoco basins ( Fig. 74View FIGURE 74). Brycon amazonicus  is a typical lowland, muddy-water species. Its distribution in the Amazon basin encompasses the middle and upper section of the Rio Amazonas /Solimões, from Pucallpa (8°23’S, 74°31’W) at the Río Ucayali in Peru eastward to Porto de Moz at the lower rio Xingu (1°45’S, 52°14’W) in Brazil, and the rio Madeira up to its upstream reaches at Bolivia and Brazil. The species occurs in clear-water rivers as the rio Tapajós and rio Xingu only at their lower sections, below the major rapids. At the rio Negro, the species is also confined to its lower section, migrating dowstream into the rio Amazonas /Solimões to spawn ( Borges, 1986). At the Rio Orinoco basin, Brycon amazonicus  is widespread at the middle/lower sections in both muddy- and clear-water tributaries. Brycon amazonicus  is known from the northwestern portion of Guyana, near the border with Venezuela, at the Amacuro River drainage, a river system connected with the Orinoco delta. The species is recorded from the island of Trinidad, an odd occurrence which is discussed at the “Biogeography” section. As some other highlyfecund, total spawners characiforms as Colossoma macropomum  ( Serrasalmidae  ), Brycon amazonicus  is only rarely found in the lower Amazon, apparently because the circadian variations of water level due to tidal influence does not allow the establishment of a population of the species in that river stretch ( Lima & Ribeiro, 2011: 161). Brycon amazonicus  has been extensively stocked, and escaped individuals are recorded from several river drainages in southeastern and central Brazil, but so far there is no evidence of self-sustaining, established populations outside its native range. These records are not mapped in Fig. 74View FIGURE 74.

Comparisons. Brycon amazonicus  is overall more similar, and possibly closely related to B. gouldingi  . Besides the caudal-fin color pattern, there are two additional differences between the species. One, the presence of a straight, well-defined dark stripe on the anal-fin base in B. gouldingi  (vs. a diffuse dark stripe on the anal-fin base in B. amazonicus  ), cannot, however, be used to unequivocally distinguish both species because a dark stripe is often absent in B. gouldingi  specimens larger than 200 mm SL and in some specimens of B. amazonicus  of all sizes. The second distinguishing character is lateral line counts, which present a large overlap, but are generally lower in Brycon amazonicus  (vs. 57–75, modally 68 in B. amazonicus  , vs. 66–82, modally 74, in B. gouldingi  ).

There is a striking resemblance between specimens of Brycon amazonicus  and B. hilarii  from the Peruvian portion of the Amazon basin (from Iquitos, Depto. Loreto, upriver to Pucallpa, Depto. Ucayali). Most specimens of Brycon amazonicus  from this portion of the basin possess a color pattern reminiscent of B. hilarii  , with dark pigmentantion concentrated on the middle/distal portion of the caudal peduncle and middle caudal-fin rays (see item “Variation”, above). Though the majority of the specimens of Brycon amazonicus  from the Peruvian portion of the Amazon basin present this color pattern, a few specimens from the area near Iquitos display the “typical” color pattern (e.g., ANSP 178374, MUSM 7020). Brycon amazonicus  from the Peruvian portion of the Amazon basin are not easily distinguished from the sympatric B. hilarii  , though the latter are typically more slender fish, with less angled predorsal and pre-pelvic body profiles, with clear-colored pectoral and pelvic fins (pectoral and pelvic fins typically darkened in B. amazonicus  ), and with a more developed caudal peduncle/caudal fin stripe (which in B. amazonicus  is more blotch-like). Both species are sympatric and often collected together along the Río Amazonas and Río Ucayali from Iquitos to Pucallpa. There are no records of Brycon hilarii  downstream from the Iquitos area, and no records of Brycon amazonicus  upstream from the Pucallpa area. See the discussion under the item “Putative examples of mimicry involving Brycon  ”, below.

Ecological notes. Detailed information on the ecology of Brycon amazonicus  were provided by Goulding (1979, 1980; as Brycon  sp.), and Borges (1986; as Brycon cf. erythropterum  ). Goulding (1979, 1980) studied the species at the rio Machado (rio Madeira basin, Rondônia). Goulding (1980) considered Brycon amazonicus  as a primarily seed/fruit eater during the flood season. Most importantat seed/fruits species found in guts of Brycon amazonicus  specimens caught in the flooded forest of the rio Machado were Hevea spruceana  ( Euphorbiaceae  ), Hevea brasiliensis  ( Euphorbiaceae  ), Luffa  sp. (Curcubitaceae), Neobalatia sp. ( Sapotaceae  ), and Calyptranthes ruizana  ( Myrtaceae  ) ( Goulding, 1980). Also found were terrestrial arthropods, leaves and rodent remains. Specimens caught during the dry season generally possessed empty guts ( Goulding, 1979, 1980). Complex migratory patterns are reported between the rio Madeira and clearwater tributaries, which can be summarized as a migration into the flooded forests during the early flood season, a downstream spawning migration from the clearwater tributaries into the muddy-water rio Madeira during the middle of the flood season, return to the flooded forests after the spawning, and a massive dispersive migratory movement in the early dry season, when the schools of Brycon amazonicus  may contain hundreds to thousands of individuals ( Goulding, 1979).

Borges (1986) studied diet and migrations of the species in rio Negro. Adult specimens (260–455 mm SL) presented a mean stomach fulness higher during the flood season. Fruits, seeds and arthropods were the main dietary itens, with fruits and seeds more common during the flood season and arthropods more expressive during the dry season. Among the fruits/seeds, Euphorbiaceae  ( Hevea spruceana  , Mabea caudata  , and Alchornea schomburgkiana  ) were dominant in guts contents, followed by Lauraceae  , “Leguminosae”, Arecaceae  , Moraceae  , Melastomataceae  , and Annonaceae  ( Pseudoxandra polypheba  ). Borges (1986: 50–51) noticed that the fruit/seed species that dominated in the diet were generally found crushed in stomach contents, while species presenting a relative smaller importance were generally found intact. Fishes and reptiles (lizards and snakes) were also found in low frequences in gut contents. Juvenile specimens (smaller than 185 mm SL) ingested a considerably larger proportion of arthropods and fishes when compared with adult specimens. Similarly to the observations of Goulding (1979, 1980), specimens collected in migratory schools or terra firme streams during the dry season generally possessed empty stomachs. Borges (1986: 123–125) summarized the available natural history information on Brycon amazonicus  at the rio Negro basin. This author hypothesized that Brycon amazonicus  does not spawn in this river system, since juvenile specimens are never collected in the area (nor larvae; Lima & Araújo- Lima, 2004), but are very common in the floodplains lakes of the rio Solimões/ Amazonas. Also, downstream migrations are observed in the rio Negro during the flood season (December/January), implying that in fact the species spawn in the rio Solimões/ Amazonas. A few other characiform fishes that possess similar migratory/ spawning habits are Colossoma macropomum  , Semaprochilodus taeniurus  , and S. insignis  ( Ribeiro & Petrere, 1990; Araújo-Lima & Goulding, 1997; Goulding et al., 1988), and in fact Brycon amazonicus  was reported to be associated with both Semaprochilodus  species in multi-species schools during its downstream migration ( Araújo & Ruffino, 2003), though most often they migrate in their own monospecific schools (M.C.L. Ribeiro, pers. comm..). This type of migratory behavior probably evolved as a response to the extreme oligotrophic water of the rio Negro basin, which does not allow an effective recruitment of larvae and juveniles of these highly-fecund species ( Borges, 1986; Araújo-Lima & Ruffino, 2003; Lima & Araújo-Lima, 2004). Interestingly, all the aforementioned species, as well as Brycon amazonicus  , only occur in the lower section of the rio Negro basin (see Araújo-Lima & Goulding, 1997; Castro & Vari, 2004). Similarly to the findings of Goulding (1979, 1980) in the rio Madeira basin, Borges (1986) noticed that Brycon amazonicus  return to the flooded forests of the lower rio Negro after spawning in February and March, primarily to eat seeds and fruits, and leave them for the terra firme large streams or the channel of the nearby muddy/whitewater river system (in that case, the rio Solimões/ Amazonas) during the the beginning of low water season (May to August) ( Borges, 1986; Araújo-Lima & Ruffino, 2003). Though in fact juvenile specimens of Brycon amazonicus  are abundant in the floodplains of the rio Solimões/ Amazonas (e.g., Bayley, 1988, as B. melanopterus  ; Petry et al., 2003, as B. cephalus  ), contrary to the view by Borges (1986) and Lima & Araújo-Lima (2004), they are not completely absent from the rio Negro, since a few lots of juveniles have been located in collections ( INPA 11072, MZUSP 60311, USNM 307069), indicating that a limited recruitment may occur at this system as well. Zaniboni Filho & Resende (1988, as B. cephalus  ) analysed with light microscopy the gonadal development in the species using specimens collected in the lower rio Negro and middle rio Solimões/ Amazonas and concluded that the species is, as expected, a total spawner, and that specimens smaller than 160 mm SL cannot be identified to sex by a macrocospic examination of gonads. Females are estimated to reach maturity at 3 years of age and 28 cm SL ( Arias, 2006). The species is reported to reach 50 cm TL in the rio Madeira basin ( Goulding, 1980: 68), and the largest specimen examined by Santos Filho & Batista (2009) during their broad survey of Brycon amazonicus  specimens landed at the Manaus market measured 52 cm TL. Brycon amazonicus  is well-known for the hability in developing a transitory dermal lip protuberance during periods of hypoxia ( Braum, 1983a, b; as Brycon cf. melanopterum  ), an adaptation in fact shared with some other characids (Winemiller, 1989b). An increase in number of maxillary teeth and gill rakers is reported to occur during the ontogeny of the species (Zaniboni Filho & Resende, 1988).

The species presents a great importance in fisheries across its entire range (see Barthem, 1999; Araújo-Lima & Ruffino, 2003; Santos et al., 2006) and has been one of most aquacultured fish species both in Brazil (e.g., Werder & Saint-Paul, 1979; Mendonça, 1994) and Colombia ( Arias, 2006).

Remarks. Chalceus amazonicus  was described by Agassiz (in Spix & Agassiz, 1829: 68–69) for the “fluvio Amazonum” (= rio Amazonas, Brazil). The species is named as Characinus amazonicus  at its depiction (plate 35), which is supposedly how Spix intended to name it, since he only supervised the execution of the plates ( Kottelat, 1988). The genus Characinus  was never described, and consequently constitutes a nomen nudum. Chalceus amazonicus  was considered in the literature as a synonym of Brycon opalinus  by Valenciennes (in Cuvier & Valenciennnes, 1850), and that practice was followed subsequently ( Günther, 1864; Eigenmann, 1910; Fowler, 1950), until Howes (1982: 9) removed it from that synonymy. Howes (1982: 9) considered that the description of Chalceus amazonicus  by Agassiz (in Spix & Agassiz, 1829) was very similar to the syntypes of C.carpophagus  , but he preferred to consider the name as a nomen dubium due the precariousness of the original description. Howes (1982: 9) erroneously inferred that the species was based on an iconotype, when actually the holotype was deposited at Munich (“In the Museum at Munich there is a fine specimen, 10 inches long, preserved in spirits of wine”; translation of the original description by Pethiyagoda & Kottelat, 1998: 155). Kottelat (1988) was not able to find the holotype of Chalceus amazonicus  among the material that Agassiz moved to the Musée d’Histoire Naturelle de Neuchâtel, and inferred that as most of Spix’s Brazilian material, it was very likely destroyed during the British bombing of the Bayerische Akademie der Wissenschaften during the night of 24–25 April 1944.

The original description of Chalceus amazonicus  only provides very general information on the species, which does not allow a clear association of the name to any of the known Brycon  species from the Amazon basin. The illustration of the holotype shows a relatively elongated fish, with an olivaceous dorsal coloration, a brown body and a grayish head. Counts of the illustrated holotype are 62 lateral-line scales, 9 scales rows between dorsal-fin origin and lateral-line, 7 horizontal scales between the lateral line and the pelvic-fin insertion, and 17 branched anal-fin rays. However, as noticed by Howes (1982: 9), there is a discrepancy between the number of anal-fin rays in the illustration and the number given in the description (iii, 24). The rather vague type-locality “fluvio Amazonum” indicate that the type-specimen may have been collected across a vast expanse of the Amazon basin. The expedition leaded by J. B. Spix and C. F. Martius crossed the whole extension of the rio Amazonas /Solimões in Brazil and moved upstream into the lower course of some of its tributaries, namely, the rio Negro, rio Tapajós, rio Madeira, and rio Japurá ( Papavero, 1971: map 7). However, in the diary of the expedition, Spix & Martius (1981: 96) reported that Chalceus amazonicus  was collected at the rio Amazonas, a little above the canal de Uruará, which is situated slightly below Monte Alegre, Pará state. It is therefore assumed that the lower Amazon should be considered the area of provenance of the holotype of Chalceus amazonicus  . Two Brycon  species are commonly found in the lower rio Amazonas channel and floodplains. One of them is Brycon melanopterus  , a species diagnosed by the presence of a conspicuous oblique dark stripe extending from pelvic-fin basis to the upper lobe of caudal fin (see under this species, below). As remarked above, the plate of Chalceus amazonicus in Spix & Agassiz (1829)  shows a specimen with a uniform color pattern, without blotches or stripes. The description by Agassiz is also uninformative in regard to any dark pigmentary feature that might eventually be present in the specimen (“head bluish, but the rest of the entire fish is olive grey and bright gold”; Pethiyagoda & Kottelat, 1998). Since the oblique stripe in Brycon melanopterus  is conspicuous even in highly-faded specimens, it seems unlikely that Chalceus amazonicus  corresponds to this species. It is more likely that Chalceus amazonicus  refers to a species described subsequently several times under different names, beginning with Valenciennes (in Cuvier & Valenciennes, 1850) as Chalceus carpophaga  . Since the holotype of Chalceus amazonicus  was lost, a neotype need to be designated in order to dispel any doubt concerning the identity of the species. Therefore, we herein designate the specimen INPA 3415, collected in the rio Trombetas, a tributary of the Rio Amazonas at Pará state, a locality relatively close to Monte Alegre, as the neotype of Chalceus amazonicus Agassiz. 

Brycon amazonicus  , as herein recognized, comprises seven synonyms. Valenciennes (in Cuvier & Valenciennes, 1850: 252–253) described Chalceus carpophaga  from the “Essequibo” and “Amazone”. In spite of the fact that Valenciennes have not selected a holotype for the species, the description of the species is clearly based on a specimen said as have being collected at the Essequibo River by Schomburgk. Bertin (1948: 15) reported three syntypes for the species: MNHN A.9832 (“Fl. Essequibo”), MNHN 98 (Amazon River), and MNHN A.8615 (Sabará, Minas Gerais). Géry & Mahnert (1992: 816) appropriately designated the specimen MNHN A.9832 as the lectotype of the species, and noticed that while the paralectotype MNHN 98 is possibly conspecific to the lectotype, the paralectotype MNHN A.8615 is probably a specimen of Brycon orthotaenia  , a supposition confirmed in the present study (see item “Remarks” of B. orthotaenia  ). The lectotype of Chalceus carpophaga  is rather faded, but longitudinal wavy stripes can still be recognized in the specimen. This is a feature shared by some Brycon  species, among them, B. amazonicus  , which is the only species presenting this pigmentary feature known to occur in northeastern South America. Morphometric and meristic data are within the range of variation observed in Brycon amazonicus  . Chalceus carpophaga  is therefore considered as a junior synonym of Brycon amazonicus  . Though Valenciennes (in Cuvier & Valenciennes, 1850) mentions “Essequibo” as the locality for the lectotype of Chalceus carpophaga  , the label of the specimen records its locality as being merely “Guyane Anglaise”. Just a few more specimens of Brycon amazonicus  are known from Guyana (AMNH 14405 and FMNH 53353, the latter being the holotype of B. siebenthalae  , see below), all of which were collected in the Amacuro River system in northwestern Guyana (see item “Distribution”, above). Since no additional specimen of Brycon amazonicus  has ever been collected in the Essequibo River itself, we strongly suspect that the locality of the lectotype of Chalceus carpophaga  given by Valenciennes (in Cuvier & Valenciennes, 1850) is incorrect, though as a Schomburgk specimen, it was in fact very likely collected somewhere else in Guyana, more likely at the Amacuro river system (see below).

Günther (1869: 423–424) described Megalobrycon cephalus  based on two specimens ( BMNH 1869.5.21: 1–2) collected by Edward Bartlett at the “Upper Amazons”. Judging for another collections by Bartlett sent to the British Museum, these specimens were likely collected in the peruvian portion of the Amazon basin (e.g., Günther, 1868: 229; Sclater & Salvin, 1866). Günther (1869) did not compared Megalobrycon cephalus  to any Brycon  species, remarking only at the generic diagnosis that Megalobrycon  would be “closely allied to Bryconops  ” (see comments on the item “Remarks” of the generic section, above). Both syntypes are now in a very poor state of preservation, with almost no scales left on the body, fins broken, and cranial bones in process of disarticulation. In fact, these specimens were already reported to be in a poor state of preservation when first studied ( Günther, 1869). The best preserved syntype possess wavy longitudinal stripes in the scales of the upper portion of the caudal peduncle (the only portion of body with a large number of scales still present) and about 58 lateral-line scales (counts obtained through counting scale pockets). These two features combined suggests that the syntypes of Brycon cephalus  refer to B. amazonicus  , since the other Brycon  species with wavy longitudinal stripes known to occur in the upper Amazon basin in Peru, B. hilarii  , possess considerably higher lateral-line scales counts (67–82). We thus consider Brycon cephalus  as a synonym of B. amazonicus  . Brycon cephalus  was until the recent past the name usually employed for B. amazonicus  in the Central Amazon basin (e.g., Regan, 1905; Howes, 1982; Zaniboni Filho & Resende, 1988; Géry & Mahnert, 1992; Ferreira et al., 1998; Araújo-Lima & Ruffino, 2003). Howes (1982), Zaniboni Filho & Resende (1988), and particularly Géry & Mahnert (1992) studied in detail the variation and diagnostic features of Brycon amazonicus  , under the name B. cephalus  , comparing it with the syntopic B. melanopterus  .

Cope (1872: 261–262) described Brycon capito  from “Ambyiacu” (= Río Ampyiacu, near Pebas, Loreto, Peru), the same type-locality of Megalobrycon erythropterum  (a synonym of B. hilarii  , see item “Remarks” of the latter species). Cope (1872) did not compared Brycon capito  with any other Brycon  species, nor discussed why he considered it to be a new species. The holotype of Brycon capito  ( ANSP 8058) was examined during the course of the present study, and is a juvenile Brycon  specimen in very poor condition of preservation, being desiccated, scaleless, and faded ( Fig. 68View FIGURE 68). Lateral-line counts of the holotype of Brycon capito  were low, according to Cope (1872) (56 lateral line scales), a count considerably lower than counts of the sympatric B. hilarii  (67–82) but very close to the range of B. amazonicus  (57–75). The holotype of Brycon capito  does not seem to be distinct from B. amazonicus  juveniles (compare Figs. 70View FIGURE 70 and 75View FIGURE 75), and thus, we consider the earlier as a synonym of the latter.

Steindachner (1879a: 150) briefly described Brycon longiceps  , without mentioning its type-locality. A little later, a proper description of the species was published ( Steindachner, 1879b: 156–157). Steindachner (1879b) described the species based on three juvenile specimens (NMW 62944: 1–3), collected in the Río Orinoco at Ciudad Bolívar, Venezuela. The examination of the three syntypes revealed them to represent juveniles of Brycon amazonicus  . Interestingly, Steindachner (1879b) suggested that the species was in fact more similar to Brycon carpophagus  (= B. amazonicus  ), and that he was precluded of doing an adequate comparison between both species because the type series of B. longiceps  consisted in juvenile specimens, while B. carpophagus  was solely known from a few large and poorly preserved specimens.

Eigenmann (1912: 372) described Brycon siebenthalae  from a single specimen collected at “Mud Creek, Aruka River”, an independent coastal drainage from northwestern Guyana, near the border with Venezuela. We examined the holotype of the species and concluded it to be a synonym of Brycon amazonicus  , a conclusion actually earlier advanced by Géry & Mahnert (1992: 816:“présente tous le caractères de B. carpophagus  … il s’agit très probablement d’un synonym”). As remarked above, there are few Brycon amazonicus  specimens reported from Guyana, and the species appears to be restricted in that country to the northwestern river basins, which are connected to the Orinoco delta via the Amacuro River (see item “Distribution”, above).

Holly (1929: 208–209) described Brycon pellegrini  from Manaus, in the lower rio Negro /central Amazon area, Brazil. Holly (1929) does not compared the species to any congener, only mentioning that it was probably related to Brycon moorei  . Both Howes (1982: 41) and Géry & Mahnert (1992: 797) considered this species to be a junior synonym of B. cephalus  (= Brycon amazonicus  ). The holotype of Brycon pellegrini  was not examined during the present study, but its original description provides enough information to allow a safe judgement concerning its taxonomic status. At the Manaus area, four Brycon  are known: B. pesu  , B. falcatus  , B. melanopterus  , and B. amazonicus  . Holly (1929) mentions that the holotype of Brycon pellegrini  possess the caudal-fin basis darkened (“Schwanzflossenbasis ist dunkler”). The only Brycon  species occurring in the Manaus area to conform such a description of the caudal-fin basis is B. amazonicus  , since the remaining species either lack dark pigmentation at the basis of caudal fin ( B. pesu  ) or possess a very characteristic caudal-basis color pattern that would not be so succinctly described if present ( B. falcatus  and B. melanopterus  ). In addition, the stated scale counts for the holotype of Brycon pellegrini  , 69/14/9 ( Holly, 1929) are beyond the upper limits known for B. falcatus  and B. pesu  , but within the range of both B. melanopterus  and B. amazonicus  . Finally, Holly (1929: 209) mentions that the fins are brownish (“Die flosse sind braun”). Darkened paired fins are found, among the Brycon  species present in the Manaus area, only in B. amazonicus  . Consequently, and confirming the view advanced by Howes (1982) and Géry & Mahnert (1992), there is enough evidence to consider Brycon pellegrini  as a synonym of B. amazonicus  .

Brycon siebenthalae iquitensis Nakashima (1941: 69–70)  was described having as its type-locality “las cercanías del puerto de Iquitos”. The type material was apparently never deposited in a fish collection and is lost (H. Ortega, pers. com.). Nakashima (1941) did not provided any reasoning about why to describe a subspecies of Brycon siebenthalae  from the Amazonian portion of Peru. Presumably, he merely gave a subspecific status to a Brycon  population he considered to be conspecific with Brycon siebenthalae  from Guyana. Though the inaccurate description given by Nakashima (1941), compounded with the lack of type-material precludes a definitive conclusion, we consider that the best alternative available to merely consider Brycon siebenthalae iquitensis  as a synonym of B. amazonicus  .

Both Kner (1860) and Amaral-Campos (1950: 140) mixed in their accounts of Brycon hilarii  specimens authentically belonging to the latter species (from “Cujaba” and “ Mato Grosso ”, respectively) with specimens of B. amazonicus  , probably because of the high scales counts shared by both species. Kner (1860: 11) cites a specimen collected by J. Natterer at “ Salto Theotonio” (now spelled Salto Teotônio, at the rio Madeira, Rondônia, Brazil), under the common name “Schatö-arana”, an obvious mispelling of “jatuarana”, the common name by which Brycon amazonicus  is still known in the rio Madeira basin.

Braum (1983a, b) studied the ability of a Brycon  species from central Amazon in developing oral dermal protuberances to withstand hypoxia. Although he identified the species he studied as Brycon cf. melanopterus  , a picture published in one of the articles ( Braum, 1983a: fig. 1a) shows that, instead, that he has studied B. amazonicus  . Lasso (1992: 11) identified Brycon melanopterus  from the lower río Suapure, Río Orinoco basin. The picture presented in the paper (fig. 3, p. 22) shows, instead, a specimen of Brycon amazonicus  . Brycon melanopterus  is in fact restricted to the Amazon basin (see item “Distribution” and “Comparisons” of this latter species). Bayley (1988) studied the growth of several young fish species occurring in the floodplains of rio Solimões near Manaus, including what that author has identified as Brycon melanopterus  . Examination of the material collected by Bayley in both USNM and MZUSP collections showed that, although there are in fact a few Brycon melanopterus  specimens in his Brycon  samples, the bulk of it is actually constituted by B. amazonicus  . Another possible misidentifications involving the species (e.g., Brycon cf. melanopterus  ; Junk et al., 1983; Brycon hilarii  ; Marlier, 1968: 56) could not be checked due to lack of information on the material which based them, but are with great likelihood attributable to B. amazonicus  .

Galvis et al. (2006: 188–189) identified Brycon cephalus  (= B. amazonicus  ) from Leticia, Colombia. Though the characters used in their description allow the identification of the material examined by them as Brycon amazonicus  (e.g., the pigmentary features of the scales; Galvis et al., 2006: 189, fig. 85d), the picture of the species (p. 457) depicts instead a specimen of B. melanopterus  .

Material examined. Type material: INPA 3415View Materials (1, 362.0 mm SL): Brazil, Pará, rio Trombetas, igarapé Caxipacoré , c. 0°34’S, 56°45’W; E. Ferreira & M. Jégu, 20 April 1985GoogleMaps  . Neotype of Chalceus amazonicus Agassiz, in Spix & Agassiz, 1829  , by present designation. MNHN A.9832 (1, 267.8 mm SL): “ Guyane Anglaise”; Schomburgk , no date  . Lectotype of Chalceus carpophaga Valenciennes  (designated by Géry & Mahnert, 1992: 816). MNHN 98View Materials (1, 260.9 mm SL): “ Amazone ”; Montravel , no date  . Paralectotype of Chalceus carpophaga Valenciennes  (designated by Géry & Mahnert, 1992: 816). BMNH 1869.5.21: 1 (2, 235.7–307.0 mm SL): “ Upper Amazons ”; E. Bartlett, no date  ; syntypes of Megalobrycon cephalus Günther.  ANSP 8058View Materials (1, 46.0 mm SL): “ Ecuador, Ambyiacu River ” [= Peru, Depto. Loreto, Rio Ampyiacu, trib. Río Amazonas , c. 3°21’S, 71°48’W]; J. Hauxwell, no dateGoogleMaps  . Holotype of Brycon capito Cope.  NMW 62944View Materials (49.1–67.7 mm SL): “ Orinoco bei Ciudad Bolivar ” [ Venezuela, Estado Bolivar, 8°8’N, 63°34’W]GoogleMaps  ; Schilling, 1879; syntypes of Brycon longiceps Steindachner.  FMNH 53353View Materials (1, 169.1 mm SL): Guyana, Mud Creek in Aruka River [c. 8°13’N, 59°44’W]; S.E. Shideler, 1908GoogleMaps  . Holotype of Brycon siebenthalae Eigenmann. 

Non types. Brazil, Pará state, rio Amazonas basin: MCZ 21134View Materials (1, 174.7 mm SL)  ; MCZ 21089View Materials (3, 161.4– 174.0 mm SL): Rio Xingu  , Porto de Moz, 1°45’S, 52°14’W; Thayer Expedition, Aug –Sept 1865GoogleMaps  . BMNH 1926.10.27.7 (1, 190.4 mm SL): Monte Alegre , Rio Amazonas, 2°0’S, 54°4’W; C. Ternetz, no dateGoogleMaps  . MZUSP 9501View Materials (1, 121.6 mm SL): Monte Alegre , rio Amazonas, 2°0’S, 54°4’W; EPA, 14 Aug 1968GoogleMaps  . MNRJ 11263View Materials (1, 200.8 mm SL): rio Amazonas, “ Maicura ” (probably rio Maicuru , c. 2°9’S, 54°12’W); L. Travassos & J. de Freitas, Feb 1960GoogleMaps  . MZUSP 9177 (1, 185.0 mm SL): Santarém, rio Maicá , c. 2°27’S, 54°40’W; EPA, 19–27 Oct 1971GoogleMaps  . CAS 68830View Materials (1, 159.0 mm SL): “ Lagoa Grande into Rio Amazona at Castello Branco ” (probably Lagoa Grande de Curuaí , c. 2°14’S, 55°16’W); C. Ternetz, 14 Jul 1923GoogleMaps  . CAS 68884View Materials (1, 174.7 mm SL): “ Lagoa Grande into Rio Amazon” (probably Lagoa Grande de Curuaí , c. 2°14’S, 55°16’W); C. Ternetz, Nov 1924GoogleMaps  . ZUEC 8033View Materials (1, 223.0 mm SL), Curuá, Lago Preto, fazenda São Luís (Costa do Cruzeiro, rio Amazonas), 2°6'22''S, 55° 10'17''W; F.C. TGoogleMaps  . Lima, W.G.R. Crampton, J.S. Ready et al., 22 Nov 2013. MCZ 21099View Materials (2, 139.2– 170.7 mm SL)  ; MCZ 21098View Materials (1, 155.7 mm SL): Rio Amazonas, Óbidos , 1°52’S, 55°30’W; Thayer Expedition, Nov –Dec 1865GoogleMaps  . MPEG 11181View Materials (1, 190.6 mm SL): Juruti, Igarapé Guaraná , 2°29’19’’S, 56°14’19’’ W; A. Hercos & V. Sena, 21 Aug 2006GoogleMaps  . CAS 66852View Materials (1, 94.3 mm SL): Juruti , 2°9’S, 56°6’W; no collector specifiedGoogleMaps  . ANSP 82285View Materials (1, 207.0 mm SL): “ Para ”; J.L. Troemner, no date  . Pará state, rio Tapajós basin: MZUSP 8405View Materials (3, 146.6– 156.6 mm SL)  : Santarém, igarapé Jacundá, Alter do Chão , c. 2°30’S, 54°58’W; EPA, 23 Dec 1967GoogleMaps  . MZUSP 57475View Materials (1, 258.1 mm SL): Santarém, Alter do Chão , 2°28'5''S 54°55'34''W; M. Westneat et al., 31 Oct 1994GoogleMaps  . MZUSP 60323View Materials (1, 154.5 mm SL): Santarém, rio Tapajós , 2°25’S, 54°45’WGoogleMaps  ; A. Campos, Oct 1944. FMNH 92083View Materials (1, 159.6 mm SL)  : Santarém, rio Tapajós , 2°25’S, 54°45’W; J.D. Haseman, 12 Dec 1909. MCZ 20780View Materials (1, 117.7 mm SL)GoogleMaps  ; MCZ 21094View Materials (1, 176.3 mm SL): Santarém, rio Tapajós , 2°25’S, 54°45’W; D. Bourget, 1865GoogleMaps  . MCZ 21125View Materials (3, 258.0–315.0 mm SL): Rio Tapajós, probably at its mouth; N. Dexter et al., 26 Aug 1865  . CAS 68858 (1, 150.2 mm SL); CAS 68854View Materials (1, 128.4 mm SL): market at Santarém; C. Ternetz, Aug –Sept 1924  . CAS 68837 (1, 243 mm SL); CAS 68915View Materials (3, 115.7– 121.3 mm SL): rio Tapajós , Santarém, 2°25’S, 54°45’W; C. Ternetz, July 1924GoogleMaps  . MNHN 1909-070View Materials (1, 115.1 mm SL): Santarém, 2°25’S, 54°45’W; C. Jobert, no dateGoogleMaps  . Pará state, rio Trombetas basin: MZUSP 8255View Materials (1, 151.0 mm SL): rio Trombetas, Oriximiná , 1°47’S, 55°52’W; EPA, 16–18 Dec 1967GoogleMaps  . MZUSP 56777View Materials (3, 207.2– 231.6 mm SL): rio Trombetas, Cuminá , c. 1°31’S, 56°2’W; M. Goulding, Oct –Nov 1983GoogleMaps  . INPA 16449View Materials (1, 341.3 mm SL): rio Cachorro (trib. rio Trombetas ), c. 1°0’S, 57°3’W; E. Ferreira & L.H.R. Py-Daniel, 18 Oct 1985GoogleMaps  . INPA 16440View Materials (1, 364.1 mm SL): rio Trombetas, near the mouth of igarapé Caxipacoré , c. 0°35’S, 56°47’W; E. Ferreira & L.H.R. Py-Daniel, 16 Oct 1985GoogleMaps  . INPA 3416View Materials (1, 317.6 mm SL): rio Mapuera (trib. rio Trombetas), Cachoeira da Égua , c. 1°5’S, 57°20’W; E. Ferreira, 17 May 1985GoogleMaps  . Amazonas state, rio Solimões / Amazonas basin: MZUSP 5801View Materials (2, 317.2– 329.7 mm SL): Lago Saracá, Silves  , c. 2°53’S, 58°21’W; EPA, 17–18 March 1967GoogleMaps  . MCZ 21124View Materials (1, 176.5 mm SL): Lago Saracá at Silves , c. 2°53’S, 58°21’W; S.V.R. Thayer, Dec 1865GoogleMaps  . MCZ 21086View Materials (2, 147.0– 183.2 mm SL): Rio Amazonas, Parintins , 2°38’S, 56°45’W; L. Agassiz, 27 Aug –2 Sept 1865GoogleMaps  . MCZ 21091View Materials (1, 160.4 mm SL): Lago José Assú [= Igarapé Açu or rio Andirá , 2°54’S, 57°8’W]; L. Agassiz, 27–30 Aug 1865GoogleMaps  . MZUSP 13430View Materials (1, 323.2 mm SL): Itacoatiara , rio Amazonas, 3°9’S, 58°27’W; N. Smith, 28 Sept 1977GoogleMaps  . MNHN 1996-1085 (1, 184. mm SL); MNHN 1996-1089 (1, 191.8 mm SL); MNHN 1996-1081 (1, 291.6 mm SL); MNHN 1996-1082 (1, 215.1 mm SL); MNHN 1996-1083 (1, 218.3 mm SL); MNHN 1996-1087View Materials (1, 171.5 mm SL): rio Urubu   , Salto Lindóia, c. 2°37’S, 59°22’W; M. Jégu, Sept –Oct 1993GoogleMaps  . MCZ 92873View Materials (1, 102.7 mm SL): Lago Jacaretinga, rio Amazonas, near Manaus; T.J. Zaret et al., 9 Jan 1980  . INPA 16384 (2, 88.5–101.6 mm SL); INPA 16450View Materials (3, 143.4– 293.8 mm SL): rio Amazonas, Ilha do Careiro, Lago do Rei, c  . 3°9’S, 59°47’W; Eq. Ictiologia/INPA, June –Oct 1986. USNM 307069 (1, 125.3 mm SL); Rio Negro, Lago Janauari , c. 3°13’S, 60°1’W; P. Bayley, 13 Oct 1977GoogleMaps  . INPA 4599View Materials (2, 228.7– 241.4 mm SL): rio Amazonas, Lago do Careiro , Ilha do Rei, c. 3°9’S, 59°47’W; M. Jégu et al., 24 Feb 1986GoogleMaps  . INPA 5784 (16, 104.7– 161.9 mm SL); INPA 5785 (8, 65.9–77.5 mm SL); INPA 5787 (14, 99.9– 163.9 mm SL); INPA 11071 (18, 49.1–107.7 mm SL); MZUSP 60310 (4, 53.0– 95.1 mm SL); INPA 16348 (1, 44.8 mm SL); INPA 16390 (119, 27.2–77.1 mm SL); INPA 16432 (59, 42.5–99.0 mm SL); INPA 16430 (29, 34.0– 51.1 mm SL); INPA 16455 (3, 38.2–48.9 mm SL); INPA 16454 (1, 112.9 mm SL); INPA 16444View Materials (1, 44.5 mm SL): rio Solimões , Ilha da Marchantaria, c  . 3°14’S, 59°55’W; Eq. Ictiologia/INPA, 1976–1981. INPA 13323 (11, 46.0– 93.7 mm SL); INPA 13346 (9, 41.1–66.6 mm SL); INPA 13315 (15, 38.0–74.0 mm SL); INPA 13311 (28, 30.3–82.7 mm SL); INPA 13320 (43, 14.2–60.9 mm SL); INPA 13316 (2, 42.5–43.4 mm SL); INPA 13319 (14, 36.8–68.1 mm SL); INPA 13306 (1, 59.9 mm SL); INPA 13314 (1, 100.4 mm SL); INPA 13319 (7, 47.6–53.6 mm SL); INPA 13300 (5, 43.6–83.9 mm SL); INPA 13310 (3, 50.8–57.8 mm SL); INPA 13326 (7, 61.6–70.3 mm SL); INPA 13339 (2, 64.0–71.0 mm SL); INPA 13299 (32, 26.4–49.7 mm SL); INPA 13317 (1, 39.8 mm SL); INPA 13303 (1, 60.5 mm SL); INPA 13307 (1, 25.4 mm SL); INPA 13324 (16, 31.9–70.9 mm SL); INPA 13328 (37, 12.7–37.7 mm SL); INPA 13329 (1, 62.5 mm SL); INPA 13309 (1, 34.7 mm SL); INPA 13321 (4, 39.4–57.2 mm SL); INPA 13327 (1, 64.6 mm SL); INPA 13312 (1, 37.6 mm SL); INPA 13301 (2, 28.8–40.3 mm SL); INPA 13302 (2, 16.9– 32.7 mm SL); INPA 13318View Materials (7, 40.0–65.0 mm SL): rio Solimões , Ilha da Marchantaria, c. 3°14’S, 59°55’W; P. Petry & R. Sotero, Jan –March 1993GoogleMaps  . MZUSP 75565 (3, 72.8–80.9 mm SL); MZUSP 74668 (16, 44.8–66.5 mm SL); USNM 307081 (237, 13.4–31.6 mm SL); USNM 307028 (31, 13.5–19.4 mm SL); USNM 307064 (80, 13.0– 39.0 mm SL); USNM 371013 (10, 35.1–53.7 mm SL); USNM 371014 (1, 80.8 mm SL); USNM 307065 (2, 61.5– 66.9 mm SL); USNM 307032 (5, 31.0– 82.6 mm SL); USNM 307072 (7, 57.9–86.2 mm SL); USNM 307071 (261, 16.4–66.7 mm SL); USNM 307061 (2, 158.3–162.0 mm SL); USNM 307002 (1, 110.1 mm SL); USNM 307023 (1, 73.7 mm SL); USNM 307018 (2, 75.7–84.1 mm SL); USNM 307060 (2, 159.6– 174.6 mm SL); USNM 307024View Materials (6, 27.9–86.2 mm SL): rio Solimões , Ilha de Marchantaria, c. 3°14’S, 59°55’W; P. Bayley, 1977–1979GoogleMaps  . INPA 16421 (2, 29.2–40.7 mm SL): “Lago Janauacá com Marchantaria”; Eq. Ictiologia/INPA, 19 Jan 1977. MZUSP 75567 (2, 114.6– 116.3 mm SL); MZUSP 75568 (2, 108.0–117.0 mm SL); USNM 307062 (1, 160.5 mm SL); USNM 307014 (4, 59.7–76.5 mm SL); USNM 371015 (1, 126.2 mm SL); USNM 307070 (3, 108.5–124.0 mm SL); USNM 307154 (3, 43.5–96.2 mm SL); USNM 307006 (12, 18.6–46.6 mm SL); Rio Solimões, Lago Janauacá lakes complex, c. 3°22’S, 60°12’W; P. Bayley, 1977–1979. INPA 16451View Materials (4, 37.4–54.9 mm SL)GoogleMaps  ; INPA 16364View Materials (1, 156.7 mm SL): rio Solimões , Lago Janauacá, c  . 3°24’S, 60°18’W; Eq. Ictiologia/INPA, Jan 1977. ZUEC 7823View Materials (1,159.1 mm SL): Manaquiri, Lago Janauacá , c. 3°24’S, 60°18’W; V.S. Uieda, 11–12 Aug 1979GoogleMaps  . ZUEC 5345View Materials (1, 89.9 mm SL): Manaquiri, Lago Janauacá , c. 3°24’S, 60°18’W; U. Caramaschi, July 1977GoogleMaps  . MZUSP 6627View Materials (3, 123.1– 149.5 mm SL): igarapé tributary of Lago Manacapuru , c. 3°17’S  . 60°54’W; EPA, 13 Nov 1967. MZUSP 5903View Materials (3, 84.5–176.4 mm SL): Lago Jacaré, right margin of rio Solimões , above Manacapuru, c  . 3°30’S, 60°47’W; EPA, 29– 31 March 1967. BMNH 1925.10.28.90 (1, 240.0 mm SL): Manacapuru, Rio Solimões, c. 3°17’S  . 60°54’W; Ehrhardt, no date. MZUSP 6306View Materials (1, 164.6 mm SL): Lago Castro, rio Purus at its mouth, c  . 3°43’S, 61°27’W; EPA, 7–8 Nov 1967. MZUSP 5990View Materials (1, 70.5 mm SL): rio Purus , at its mouth, c. 3°43’S, 61°27’W; EPA, 1–5 May 1967GoogleMaps  . INPA 17122View Materials (1, 207.0 mm SL): rio Purus, mouth of Paraná do Jarí, near lago Arunã , 4°55’33’’S, 62°21’58’’W; L. Rapp Py-Daniel et al., 9 June 2001GoogleMaps  . INPA 19106 (1, 178.0 mm SL); MCP 29759View Materials (1, 167.5 mm SL): Maraã, Lago Amanã, mouth of igarapé Uxi , 2°32’42’’S, 64°40’7’’W; W.G.R. Crampton, 14 Dec 1997GoogleMaps  . MCP 29757View Materials (1, 162.1 mm SL): Maraã, Lago Amanã, mouth of rio Baré , 2°27’23’’S, 64°43’35’’W; W.G.R. Crampton, 17 Dec 1997GoogleMaps  . MZUSP 27924View Materials (2, 122.9– 165.6 mm SL): Lago Mamirauá, mouth of rio Japurá , c. 2°59’S, 64°56’W; P.E. Vanzolini, 25 Sept 1983GoogleMaps  . INPA 19108View Materials (1, 139.6 mm SL): rio Solimões , Reserva Mamirauá, Lago Mamirauá; W.G.R. Crampton, Oct 1993  . INPA 19118View Materials (1, 39.0 mm SL): rio Solimões , Reserva Mamirauá, Lago Secretaria; W.G.R. Crampton, March 2001  . INPA 19107View Materials (2, 119.6– 133.5 mm SL): rio Solimões , Reserva Mamirauá, lago Curuçá; W.G.R. Crampton, 4 Nov 1997  . MCP 29758View Materials (1, 143.1 mm SL): Alvarães, Lago Rato (channel), Lago Mamirauá system, 3° 2’58’’S, 64° 51’31’’W; W.G.R. Crampton, 11 Oct 1999GoogleMaps  . INPA 19111View Materials (1, 159.6 mm SL): Tefé, rio Solimões , Lago Capivara, Costa das Capivaras; W.G.R. Crampton, 13 Jan 2001  . MCZ 162214View Materials (1, 196.0 mm SL): Tefé , 3°24’S, 64°45’W; L. Agassiz et al., Oct 1865GoogleMaps  . INPA 19109View Materials (1, 167.1 mm SL): rio Tefé , Ilha do Martelo; W.G.R. Crampton, 14 July 1999  . MNHN 1909-0289View Materials (1, 153.5 mm SL): Rio Solimões, Tonantins , 2°52’S, 67°46’W; C. Jobert, no dateGoogleMaps  . MZUSP 55821View Materials (1, 23.4 mm SL): rio Solimões , 17.1 km downstream rio Içá, 3°0’27’’S, 67°52’46’’W; J.P. Friel et al., 22 Nov 1993GoogleMaps  . MZUSP 56153View Materials (3, 24.3–38.6 mm SL): rio Solimões , 26.1 km downstream rio Içá, 2°58’5’’S, 67°49’51’’W; J.P.Sullivan et al., 23 Nov 1993GoogleMaps  . MZUSP 27334 (2, 236.0–237.0 mm SL): Benjamin Constant, Costa do Capacete, rio Solimões , c. 4°20’S, 69°58’W; L.PGoogleMaps  . Portugal, 13 Nov 1982. MZUSP 27330View Materials (1, 184.0 mm SL): Tabatinga, Lago Caial, rio Solimões , c. 4°20’S, 69°58’W; L. PGoogleMaps  . Portugal, 8 Oct 1982. MNRJ 11262View Materials (1, 209.5 mm SL): rio Queixito, at its mouth at rio Javari , 4°20’S, 70°12’W; A. Parko, 1942GoogleMaps  . Amazonas state, rio Uatumã basin: INPA 5685View Materials (1, 258.0 mm SL): rio Uatumã, Balbina , c. 1°55’S, 59°28’W; M. Jégu, Sept 1985GoogleMaps  . INPA 15421View Materials (2, 286.1– 292.8 mm SL): rio Uatumã, Balbina , c. 1°55’S, 59°28’W; P. Chaves, July 1985GoogleMaps  . INPA 16433View Materials (2, 270.2– 284.2 mm SL): rio Uatumã , igarapé do Barreto, c  . 1°58’S, 59°27’W; Eq. Ictiologia/ INPA, 27 April 1983. INPA 5517View Materials (1, 274.7 mm SL): rio Uatumã , igarapé do Arraia; S. Amadio, May 1985  . INPA 16428View Materials (1, 287.5 mm SL): igarapé do Arraia, rio Uatumã ; Eq. Ictiologia / INPA, 15 April 1983  . INPA 16429View Materials (1, 289.4 mm SL): Amazonas, igarapé Água Branca, rio Pitinga (trib. rio Uatumã ), c. 0°52’S, 59°27’W; P. Chaves, 17 Oct 1983GoogleMaps  . INPA 16374View Materials (1, 228.9 mm SL): igarapé da Água Branca, rio Pitinga , c  . 0°52’S, 59°27’W; Eq. Ictiologia / INPA, 17 April 1983. Amazonas stateGoogleMaps  , rio Negro basin: MCZ 15995View Materials (2, 27.9–32.3 mm SL)  : Rio Negro, Lago Aleixo , c. 3°5’S, 59°53’W;. V.R. Thayer, 1865. BMNH 1943.4.24.37 (1, 196.3 mm SL): “ Manaos ”; J.C. Anthony, no dateGoogleMaps  . MZUSP 9574 (1, 178.3 mm SL): Manaus (fish market); EPA, 17–19 Sept 1968. MZUSP 6766View Materials (1, 206.0 mm SL): Manaus, igarapé Tarumãzinho , trib rio Negro, c. 3°3’S, 60°7’W; EPA, 17–18 Nov 1967GoogleMaps  . INPA 15925View Materials (1, 344.0 mm SL): rio Negro, Maruauru (not located); Cidomar, March 2000  . MZUSP 61882View Materials (5, 394.5– 412.5 mm SL): rio Puduari , upper portion, c. 2°8’S, 61°15’W; G. Borges, Nov 1981GoogleMaps  . MZUSP 105603 (14, 235.7– 428.2 mm SL): rio Negro, Praia Grande , c. 2°31’S, 69°59’W; G. Borges, 23 May 1982. INPA 2614View Materials (1, 340.9 mm SL)GoogleMaps  : rio Negro, Praia Grande , c. 2°31’S, 69°59’W; Eq. Ictiologia, 2 Jan 1984GoogleMaps  . UMMZ 217848View Materials (1, 221.5 mm SL): Arquipélago de Anavilhanas, rio Negro; W.F. Fink et al., Jan 1983  . UMMZ 217803View Materials (1, 229.4 mm SL): rio Negro, lago “ Chiclaua ” (not located); W.L. Fink, 10 Jan 1983  . INPA 9900View Materials (1, 254.7 mm SL): rio Jaú, igarapé Miratuca , c. 1°58’S, 61°30’W; M. Garcia  & Oliveira, 29 Oct 1994GoogleMaps  . INPA 11072 (20, 34.4–48.9 mm SL); MZUSP 60311 (8, 31.4–51.8 mm SL): Rio Negro, Anavilhanas , c. 2°31’S, 69°59’W; Eq. Ictiologia / INPA, 28 May 1976GoogleMaps  . INPA 16447 (13, 222.6– 316.8 mm SL): rio Negro, “ Ponta da Piraíba ” (not located); Eq. Ictiologia / INPA, 15 Aug 1985  . MNRJ 6268View Materials (1, 147.4 mm SL): rio Negro, “ Uyipiranga ” (not located)  ; A. Parko, 1941  . Amazonas state, rio Madeira basin: MZUSP 7039View Materials (11, 169.8– 269.7 mm SL): rio Canumã , c. 4°2’S, 59°6’W; EPA, 28–29 Nov 1967GoogleMaps  . MZUSP 59001View Materials (1, 270.8 mm SL): mouth of rio Madeira (purchased at the Itacoatiara fish market)  ; A.M. Zanata, 3 Aug 1996. MCZ 21090View Materials (1, 203.8 mm SL)  : Rio Madeira, probably near confluence with rio Solimões ; L. Agassiz et al., 1 Sept 1865  . Rondônia state, rio Madeira basin: MZUSP 14017View Materials (4, 301.0– 348.6 mm SL): rio Machado , at its mouth, 8°4’S, 62°53’W; M.Goulding, 18 Sept 1977GoogleMaps  . UF 100627View Materials (1, 194.0 mm SL): rio Jamari, ca. 20 km downstream from Samuel dam, locally called Pedra de Sant'anna, c  . 8°44’S, 63°29’W; J. P. Viana, 3 June 1993. INPA 16383View Materials (11, 133.0– 156.8 mm SL): rio Jamari , Samuel dam, diverting channel (cofferdam), cGoogleMaps  . 8°45’S, 63°27’W; G.M. Santos, 7–9 June 1988. INPA 16456View Materials (1, 383.4 mm SL): rio Jamari, 5 km above Samuel dam, c. 9°32’S, 63°8’W; G.M. Santos, 14 July 1985GoogleMaps  . INPA 16442View Materials (2, 282.8– 303.2 mm SL): rio Jamari, above Samuel dam, c. 9°32’S, 63°8’W; G.M. Santos, 7 Dec 1984GoogleMaps  . INPA 16443View Materials (2, 273.9– 318.5 mm SL): rio Jamari below Samuel dam, c. 8°30’, 63°29’W; G.M. Santos, 27 March 1986  . CAS 96131View Materials (1, 207.0 mm SL): Rio Madeira, “ Cachoele de Theotone ” [= Cachoeira do Teotônio , 8°52’S, 64°3’W]; J.D. Haseman, 30 Oct 1909GoogleMaps  . CAS 18395View Materials (1, 273.0 mm SL): Rio Madeira, “whirlpools at Cachoele de Girao ” [= Cachoeira do Girau , 9°17’S, 64°39’W]; J.D. Haseman, 26 Oct 1909GoogleMaps  . MZUSP 88028View Materials (1, 194.0 mm SL): rio Madeira, Estação Ecológica Antônio Mujica Nava , 9°24’50’’S, 64°56’32’’W; L.F. Silveira, 6–18 Feb 2002GoogleMaps  . MCP 39586View Materials (1, 188.4 mm SL): Igarapé Taquarás , trib. Rio Mamoré, BR-425, 9°57’43’’S, 65°17’45’’WGoogleMaps  ; A.R. Cardoso, V. Bertaco, F.C.T  . Lima & J.F. Pezzi da Silva, 25 Jul 2004. INPA 16371View Materials (1, 249.0 mm SL): rio Novo, trib. rio Guaporé , at its mouth, c  . 10°52’S, 65°16’W; G.M. Santos, 20 June 1985. INPA 16375View Materials (2, 192.7– 207.4 mm SL): Guajará-Mirim, rio Pacaás Novos (trib. rio Guaporé ), cGoogleMaps  . 11°5’S, 65°8’W; G.M. Santos, 16 Nov 1984. INPA 16377View Materials (1, 185.5 mm SL): rio Mamoré, mouth of rio Guaporé, Surpresa , 11°53’S, 65°1’W; G.M. Santos, 19 June 1984GoogleMaps  . AMNH 39956View Materials (1, 392.4 mm SL): Rio Guaporé, 5 km SW Costa Marques, c. 12°30’S, 64°17’W; R.M. Bayley et al., 23–24 Sept 1964GoogleMaps  . Mato Grosso state, rio Madeira basin: MZUSP 105602View Materials (2, 420.0–485.0 mm SL): rio Verde, trib. rio Guaporé , Brazil / Bolivia border, 14°7’36’’S, 60°28’30’’W; OGoogleMaps  . A. Cantelmo & L. Barbosa, 13–21 Aug 2005  . Bolivia, rio Madeira basin: FMNH 107242View Materials (1, 445.0 mm SL): Estado Pando  , Río Nareuda (trib. Río Orthon, Río Madre de Dios drainage), c. 11°16’S, 69°3’W; NGoogleMaps  . A. Menezes et al., 4 Sept 1996  . MNHN 1989–1430View Materials (3, 140.2– 158.8 mm SL): Estado Beni  , Río Tijamuchi (trib. Rio Beni), below Trinidad, c. 14°11’S, 65°3’W; L. Lauzanne & G. Loubens, 26–28 Apr 1984GoogleMaps  . Peru, Depto. Loreto  , Rio Ucayali /Amazonas basin:  NRM 23671View Materials (1, 113.7 mm SL); NRM 23672 (1, 123.8 mm SL): Quebrada Tocón Grande and marginal pools, at km 33 of carretera Iquitos-Nauta, Río Itaya drainage, c. 4°2’S, 73°26’W; S.O. Kullander et al., 3–4 Jul 1986GoogleMaps  . MZUSP 15266View Materials (2, 87.2–90.7 mm SL): Caño Lupuna, Río Amazonas (not located); M.V. Correa  , 10 Jun 1980  . INHS 55030View Materials (1, 118.0 mm SL): south bank trib. Río Napo, near Mazán, ca. 33 km Iquitos ,c. 3°30’S, 73°6’W; M.H. Sabaj et al., 31 Jul 1999GoogleMaps  . INHS 43857View Materials (1, 95.9 mm SL): small cocha trib. Río Napo, Isla Milagros across the town of Mázan, 3°28'59,9'’S, 73°05'05,5'’W; M.H. Sabaj et al., 2 Aug 1997  . INHS 106469View Materials (1, 134.4 mm SL): Río Nanay, Pampa Chica, 3°45'08,8'’S, 73°17'00,1'’W; M.H. Sabaj & J.W. Armbruster, 22 Jul 1997  . CAS 16031 (1, 293.7 mm SL); CAS 68880View Materials (1, 198.4 mm SL): Rio Amazonas, Iquitos; W.R. Allen, Sept 1920  . CAS 160579 (1, 132.4 mm SL); CAS 160749 (1, 120.7 mm SL); CAS 136578View Materials (1, 164.3 mm SL): Cano del Shansho, near Pebas , c. 3°21’S, 71°48’W; W.G. Scherer, Sept 1936GoogleMaps  . ANSP 178374 (3, 114.2– 162.7 mm SL): Prov. Maynas, Río Yanuyacu (trib. R. Amazonas at mouth of Cano Chincana, Emerald Forest Lodge, approx. 25 miles S. of Iquitos; M.H. Sabaj et al., 11 Aug 2001. MUSM 7020 (1, 134.5 mm SL)  : Río Pacaya, Cocha Zapote , 5°75’S, 74°25’W; H.W. Koepcke, March 1960. MUSM 18177 (1, 40.8 mm SL): San Pablo de TipishcaGoogleMaps  , Río Marañon; H. Ortega et al., 15 Apr 2001  . Depto. Ucayali  , Río Ucayali basin:  MZUSP 26219View Materials (3, 58.9–60.8 mm SL); MZUSP 26218View Materials (3, 60.7–66.1 mm SL): Río Ucayali, Shanahao, Prov. Coronel Portillo; H. Ortega, 29 Apr 1976  . MZUSP 25937 (6, 1 cs, 27.7–52.0 mm SL); AMNH 43338 (2, 66.1–73.5 mm SL); MUSM 41144 (1, 50.7 mm SL): Río Ucayali, Masisea , Prov. Coronel Portillo, 8°35’S, 74°20’W; H. Ortega, 21–23 Nov 1973GoogleMaps  . MUSM 41154 (1, 148.8 mm SL): Pucallpa, Río Ucayali, Masisea , 8°35’S, 74°20’W; H. Ortega, 24 Sept 1975GoogleMaps  . MUSM 108 (2, 133.0–134.0 mm SL): Ucayali, Pucallpa, Masisea, Lobococha , 8°35’S, 74°20’W; H. Ortega, 20 Apr 1983GoogleMaps  . MUSM 315 (2, 130.5–132.0 mm SL): Pucallpa, Río Ucayali, Utiquinía , 8°12’S, 74°34’W; H. Ortega, 20 Sept 1985GoogleMaps  . Colombia, Rio Amazonas basin: UMMZ 191053View Materials (1, 225.5 mm SL): Leticia  , Rio Amazonas, 4°13’S, 69°57’W; R. Faber, 17 May 1971GoogleMaps  . Venezuela, rio Orinoco basin, Estado Delta Amacuro: USNM 236562View Materials (50, 122.7– 297.8 mm SL): tidal stream on river shore, 49 miles from sea buoy, 8°37’36’’N, 60°49’36’’W; J.N. Baskin & D.J. Stewart, 20 Nov 1979GoogleMaps  . Estado Anzoategui: ANSP 166777View Materials (1, 172.0 mm SL): Soledad  , Laguna Curita, 8°9’30’’N, 63°35’55’’W; M. Rodriguez, 27 Apr 1988GoogleMaps  . ANSP 166480View Materials (4, 149.5– 170.9 mm SL): Soledad , Laguna Aguilera, 8°11’30’’N, 63°26’45’’W; M. Rodriguez & S. Richardson, 22 Jan 1987GoogleMaps  . Estado Guárico: MZUSP 54583View Materials (2, 142.9– 145.2 mm SL): Santa Rita  , Río Aguaro, near Santa Rita, 8°9’N, 66°14’W; J. Moscó et al., 22 Nov 1968GoogleMaps  . INHS 33942View Materials (3, 122.8– 137.7 mm SL): Pozo Azul (Río San Bartolo), P.N. Aguaro-Guariquito; W. Mejo et al., 4 Jan 1995  . INHS 34335View Materials (1, 217.5 mm SL): Río Aguaro, P.N. Aguaro-Guariquito, 15 km S Paso Cachimbo , 8°3’6’’N, 66°25’34’’W; D.C. Taphorn et al., 10 Jan 1995GoogleMaps  . FMNH 85402View Materials (6, 80.5–98.1 mm SL): borrow pit west of road, 9.7 km N of Cabruta , c. 7°43’N, 66°15’W; J.E. Thomerson et al., 5 Jan 1975GoogleMaps  . LBP 10224View Materials (4, 128.2– 148.1 mm SL): Cabruta , Río Apure, 7°37’24’’N, 66°24’48’’W; C. Oliveira & V. Tagliacollo, 21 Apr 2010GoogleMaps  . Estado Barinas: FMNH 103954View Materials (1, 154.6 mm SL): Playa Los Chicos in the Rio Suripá, ca. 2.5 hrs. above Hato Las Mercedes , c. 7°47N, 70°9’W; B. Chernoff et al., 12 Jan 1991  . Estado Apure: INHS 27660View Materials (2, 163.5– 184.4 mm SL): Caño Potrerito , hwy. bridge between San Fernando and Puerto Paez, 6° 24’48’’N, 67°31’56’’W; L.M. Page et al., 22 Jan 1992GoogleMaps  . INHS 61402View Materials (1, 178.5 mm SL): Laguna Larga (Río Cínaruco drainage), 6°33’19’’N, 67°24’49’’W; L.M. Page et al., 19 Jan 1992GoogleMaps  . ANSP 165393View Materials (1, 144.4 mm SL): Rio Claro, 15 km S of La Montaripa on San Fernando de Apure—Puerto Paez Hwy., 7°10’N, 67°25’W; S. Schaefer et al., 7 Nov 1989GoogleMaps  . ANSP 165548View Materials (3, 120.5–129.0 mm SL): flooded savannah ca. 2.0 km S of intersection of Rio Claro and San Fernando de Apure— Puerto Paez hwy., 7°10’N, 67°25’W; S. Schaefer et al., 7 Nov 1989GoogleMaps  . ANSP 165483View Materials (1, 232.0 mm SL): Río Capanaparo, backwater lagoon (mouth of caño Las Varitas ) near San Fernando de Apure—Puerto Paez hwy., 7°2’N, 67°25’W; S. Schaefer et al., 7 Nov 1989GoogleMaps  . FMNH 69900 (1, 159.3 mm SL); FMNH 117234View Materials (1, 170.0 mm SL): Río Cinaruco at confl. Madre about 60 miles S. of San Fernando de Apure, c. 6°33’N, 67°18’W; W.P. Braker & Zunwalt, Feb 1967GoogleMaps  . Estado Bolívar: ANSP 159722View Materials (1, 125.3 mm SL)  : Río Guacamayo , below bridge at crossing of Caicara-Ciudad Bolivar hwy., 7°40’N, 64°10’W; B. Chernoff & F. Provenzano, 21 Nov 1985GoogleMaps  . ANSP 159726View Materials (33, 99.1–144.4 mm SL): Caño crossing road to Las Trincheras, 2.7 km of intersection with Ciudad Bolivar-Caicara hwy., 7°22’N, 64°59’W; W. Saul et al., 20 Nov 1985GoogleMaps  . ANSP 159720View Materials (5, 104.6–220.0 mm SL): Río Agua Blanca , below bridge at crossing of Caicara-Ciudad Bolivar hwy., 7°50’N, 63°51’W; B. Chernoff et al., 21 Nov 1985GoogleMaps  . ANSP 159727View Materials (1, 117.3 mm SL): Río Caura at Maripa ferry crossing, 7°27’N, 65°12’W; B. Chernoff et al., 19 Nov 1985GoogleMaps  . ANSP 159728View Materials (2, 124.3– 194.7 mm SL): morichal Poso Vagabundo , 3 km E of Maripa, just off Caicara-Ciudad Bolivar hwy., 7°25’30’’N, 65° 9°59’W; W. Saul & R. Royero, 19 Nov 1985GoogleMaps  . ANSP 135720View Materials (1, 146.5 mm SL): Morichal Zamorai (caño), between Río Tauca and Río Tiquire , Maripa-Ciudad Bolivar hwy., 7°28’N, 64°54’W; J.E. Boehlke et al., 7 Feb 1977GoogleMaps  . ANSP 135840View Materials (2, 238.0–245.0 mm SL): mouth of small caño feeding Río Caura 15 min downstream from “hydrographic plant”, 6°20’N, 64°30’W; J.E. Boehlke et al., 22 Jan 1977GoogleMaps  . CAS uncat. (1, 152.0 mm SL); CAS 156831View Materials (3, 134.6– 167.9 mm SL): Caño de Quiribana, near Caicara , c. 7°36’N, 66°11’W; C. Ternetz, April –May 1925GoogleMaps  . CAS 154565 (1, 156.1 mm SL); CAS 154568View Materials (1, 134.9 mm SL): Rio Orinoco, mouth of Cano de Quiribana, Caicara , c. 7°36’N, 66°11’W; C. Ternetz, 5 May 1925GoogleMaps  . CAS 156758View Materials (1, 130.1 mm SL): creeks into Laguna San Raphael, Caicara , c. 7°36’N, 66°11’W; C. Ternetz, 28 Apr 1925GoogleMaps  . LBP 3026 (8, 113.7– 141.8 mm SL): Caicara del Orinoco , Río Orinoco, 7°38’12’’N, 66°19’4’’W; C. Oliveira & AGoogleMaps  . Granado, 2 Oct 2005  . ANSP 159740View Materials (14, 85.4–130.1 mm SL): Cano Caiman, at crossing of Caicara-Puerto Ayacucho hwy, 19.2 km of Ciudad Bolivar-Caicara hwy; B. Chernoff et al., 29 Nov 1985  . ANSP 159725View Materials (5, 96.7– 124.3 mm SL): River and flooded area 15 km of Maniapure on Caicara-Puerto Ayacucho hwy (Río Chaviripa?), c. 7°0’N, 66°31’W; B. Chernoff et al., 16 Nov 1985GoogleMaps  . ANSP 159724View Materials (16, 91.4–121.7 mm SL): small stream crossing Caicara-Puerto Ayacucho hwy., 18 km N of Maniapure ; B. Chernoff et al., 16 Nov 1985  . ANSP 159737View Materials (2, 100.6– 107.5 mm SL): Río Maniapure at crossing of Caicara-Puerto Ayacucho hwy., 6°55’34’’N, 66°33’32’’W; B. Chernoff et al., 28 Nov 1985GoogleMaps  . ANSP 159723View Materials (2, 18.8–110.5 mm SL): Río Caura at Puerto Las Majadas , 7°30’18’’N, 64°50’24’’W; L. Aguana, 23 Nov 1985GoogleMaps  . ANSP 159739View Materials (1, 136.2 mm SL): Caño 15.1 km E of Río Parguaza ferry crossing on Caicara-Puerto Ayacucho hwy., c. 6°19’N, 67°6’W; B. Chernoff et al., 28 Nov 1985GoogleMaps  . ANSP 159735View Materials (1, 120.3 mm SL): Cano crossing dirt road near Hacienda Fundo Malana , c. 15 km N of Sipao, 7°35’N, 65°22’W; W. Saul & R. Royero, 19 Nov 1985GoogleMaps  . ANSP 135771View Materials (1, 144.0 mm SL): Río Urbana (Urbani) on Maripa-Las Trincheras road, 7°18’N, 65°0’W; J. Boehlke et al., 20 Jan 1977GoogleMaps  . UF 80418 (2, 129.9– 140.5 mm SL): bridge of Río Chaviripa on Caicara-San Fernando do Atabapo (Puerto Ayacucho) road, c. 7°0’N, 66°31’W; D.C.GoogleMaps 

Taphorn et al., 16 Apr 1984. UF 80409 (1, 174.1 mm SL); UF 80709 (2, 131.8– 146.5 mm SL); MZUSP 62442 (2, 121.7– 126.6 mm SL): Caño Caripo/Caripito; D.C. Taphorn et al., 17 Apr 1984. FMNH 85678View Materials (1, 157.0 mm SL): Rio Orera, 13 km S. of Puerto Nuevo toward Puerto Ayacucho; J. Thomerson et al., 14 Jan 1975  . Estado Amazonas: ANSP 159738View Materials (1, 127.3 mm SL): Caño 21 km SW of El Burro , Caicara-Puerto Ayacucho hwy., c. 6°0’N, 67°21’W; W. Saul et al., 26 Nov 1985GoogleMaps  . FMNH 103953View Materials (1, 162.6 mm SL): mouth of small caño on Río Cuao, 4°58’4’’N, 67°36’55’’W; B. Chernoff et al., 12 Feb 1992GoogleMaps  . Guyana: AMNH 14405View Materials (2, 106.5– 106.9 mm SL)  ; AMNH 14323View Materials (2, 101.3– 123.9 mm SL): Santa Rosa , Amacuro River, 7°39’N, 58°57’WGoogleMaps  ; A.S. Pinkus, 1938. Trinidad: USNM 235526View Materials (1, 206.3 mm SL): Chatam stream, south west Trinidad , 10°5’N, 61°45’W; M. Atkins, 5 Sept 1980GoogleMaps  . Uncertain locality: MZUSP 3815View Materials (7, 137.1– 151.6 mm SL): “ Amazonas e Pará ”; E. Garbe, 1902. Introduced /escaped stocked specimens (all from Brazil): NUP uncat. (2, 298.5– 310.1 mm SL)  : Goiás, Caldas Novas, rio Corumbá, trib. rio Paranaíba ; Nupélia , 1996–2000. MZUSP 54008View Materials (8, 121.7– 203.6 mm SL)  : São Paulo, Pirassununga, CEPTA (stocked specimens); 27 Feb 1998  . MZUSP 82418 (6, 85.8–101.4 mm SL); MZUSP 82430View Materials (3, 111.6– 141.4 mm SL): São Paulo, Campinas, artificial pond near rio Atibaia   ; A.M. Zanata & M.P. Geraldes, March 2003  . MZUSP 86779View Materials (1, 217.5 mm SL): Rio de Janeiro, Campos, rio Paraíba do Sul (fish market); O.T. Oyakawa et al., Sept 2004  .

TABLE 18. Morphometric data of Brycon amazonicus (A: neotype of Chalceus amazonicus, INPA 3415; B: lectotype and paralectotype of Chalceus carpophagus, MNHN 9835 and MNHN 98; C: largest syntype of Brycon longiceps, NMW 62944; D; holotype of Brycon siebenthalae, FMNH 53353).

  362.0 260.9–267.8 67.7 169.1 120 67.7–485.0  
      52.6   120    
      12.6   120    
Posterior terminus of dorsal fin to hypural joint     35.6   120    
      16.1   120    
      32.9   120    
      29.1   120    
      32.3   120    
      40.8   120    
      33.2   120    

Museu de Zoologia da Universidade de Sao Paulo


Academy of Natural Sciences of Philadelphia


Instituto Nacional de Pesquisas da Amazonia


Smithsonian Institution, National Museum of Natural History


Museum National d'Histoire Naturelle


Naturhistorisches Museum, Wien


Field Museum of Natural History


Museum of Comparative Zoology


Museu Nacional/Universidade Federal de Rio de Janeiro


California Academy of Sciences


Museu de Zoologia da Universidade Estadual de Campinas


Museu Paraense Emilio Goeldi


Pontificia Universidade Catolica do Rio Grande do Sul


University of Michigan, Museum of Zoology


American Museum of Natural History


Swedish Museum of Natural History - Zoological Collections


Illinois Natural History Survey














Brycon amazonicus ( Agassiz, 1829 )

Lima, Flávio C. T. 2017

Brycon bicolor

Fernandez 2006: 54

Brycon cf. falcatus

Fernandez 2006: 55

Brycon amazonicum:

Merona 2010: 106
Leite 2004: 661

Brycon amazonicus

Lima 2013: 228
Phillip 2013: 8
Lima 2011: 151
Santos 2009: 195
Lima 2003: 175
Goulding 2003: 138

Brycon melanopterus

Lasso 1992: 11
Bayley 1988: 131

Brycon cf. cephalus

Machado-Allison 1999: 65
Almeida-Toledo 1996: 36
Ferreira 1993: 56
Goulding 1988: 124

Brycon erythropterum

Goulding 1988: 124

Brycon cf. melanopterum

Braum 1983: 355

Brycon cf. melanopterus

Piedade 2006: 1176
Werder 1984: 398
Braum 1983: 268
Werder 1983: 445
Junk 1983: 408

Brycon carpophagus:

Lima 2003: 175
Merona 2001: 387
Gery 1992: 815
Howes 1982: 15

Brycon cephalus:

Galvis 2006: 188
Galvis 2006: 457
Merona 2004: 77
Lima 2003: 253
Diaz-Sarmiento 2003: 311
Petry 2003: 565
Lima 2003: 788
Pizango-Paima 2001: 509
Ferreira 1998: 93
Gery 1992: 794
Howes 1982: 18


Fernandez 2006: 58
Piza 2002: 122
Crampton 1999: 15
Barthem 1999: 82
Goulding 1980: 68
Smith 1979: 131
Goulding 1979: 95

Brycon hilarii

Marlier 1968: 56

Brycon stubelli

Amaral 1950: 142

Brycon siebenthalae iquitensis

Nakashima 1941: 69

Chalceus taeniatus

Magalhaes 1931: 130

Brycon pellegrini

Lima 2003: 175
Howes 1982: 41
Holly 1929: 208

Brycon siebenthalae

Lima 2003: 175
Cala 1997: 55
Useche-L 1993: 341
Cala 1986: 91
Howes 1982: 44
Gery 1977: 338
Eigenmann 1912: 372

Brycon longiceps

Lima 2003: 175
Lasso 1988: 127
Mago-Leccia 1970: 69
Steindachner 1879: 150
Steindachner 1879: 157

Brycon capito

Cope 1872: 261

Megalobrycon cephalus Günther, 1869 : 423

Regan 1905: 190
Gunther 1869: 423

Chalceus hilarii

Kner 1860: 10

Chalceus carpophaga Valenciennes, in Cuvier & Valenciennes, 1850 : 252

Gery 1992: 816
Kner 1860: 12
Cuvier 1850: 252

Chalceus amazonicus Agassiz, in Spix & Agassiz, 1829 : 68

Howes 1982: 9
Spix 1829: 68