Cuvier & Valenciennes, 1850 : 246
Kner, 1860 : 10
Chalceus orbignyanus
Kner, 1860 : 11
Cope, 1872 : 263
Perugia, 1897 : 149
Boulenger, 1898b : 127
Bertoni, 1939 : 55
Géry & Mahnert, 1992 : 803
Silimon et al. , 1996 : 12
Resende et al. , 1998 : 9
Margarido & Galetti Jr., 1999 : 357
Sabino & Sazima, 1999 : 309
Britski et al. , 2007 : 42
Mateus & Estupiñan, 2002 : 165
Wantzen et al. , 2002 : 242
Resende, 2003 : 119
Lima, 2003 : 176
Brycon
hilarii
Menni, 2004 : 74
Mateus et al. , 2004 : 219
Brycon orbygnyanus
Boulenger, 1897b : 4
Ramlow, 1989 : 10
Boulenger, 1900 : 3
Eigenmann & Ogle, 1900 : 3
Fowler, 1932 : 346
Bertoni, 1939 : 55
Miranda-Ribeiro, 1940 : 44
Aguirre, 1945 : 38
Amaral Campos, 1950 : 140
Howes, 1982 : 31
Lima, 1987 : 89
Sazima & Machado, 1990 : 24
Barrella et al. , 1994 : 14
Agostinho & Júlio Jr., 1999 : 379
Lima, 2003 : 176
Makrakis et al. , 2007 : 191
Graça & Pavanelli, 2007 : 77
Sanches & Galetti, 2007 : 889
Reys et al. , 2009 : 136
Lima & Ribeiro, 2011 : 151
Bessa et al. , 2011 : 351
Salminus brasiliensis
Resende, 2011 : 471
La Monte, 1935 : 7
Marlier, 1968 : 56
Brycon hilarlii
Eigenmann & Kennedy, 1903 : 523
Fowler, 1939 : 263
Eigenmann & Allen, 1942 : 254
Ortega & Vari, 1986 : 7
Howes, 1982 : 25
Géry & Mahnert, 1984 : 176
B. melanopterus
Géry & Mahnert, 1992 : 798
Chang, 1998 : 20
Goulding et al. , 1988 : 124
Brycon cf. whitei
Saul, 1975 : 103
Ortega, 1996 : 464
A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae)
Lima, Flávio C. T.
Zootaxa
2017
4222
1
1
189
NH7H
Valenciennes, 1850
Valenciennes
1850
[151,582,836,862]
Actinopterygii
Bryconidae
Brycon
Animalia
Characiformes
97
98
Chordata
species
hilarii
Chalceus hilariiValenciennes, in Cuvier & Valenciennes, 1850: 246–248 (Type locality “Rio San Francisco”); Kner, 1860: 10– 11 (in part; “Rio Cujaba”). ? Chalceus orbignyanus(not Valenciennes): Kner, 1860: 11–12 (“ RioGuaporé”).
Salmopara-putangaKner (ex Natterer), 1860: 4 (published as a synonym of Chalceus orbygnianus; not available). Brycon melanoxanthusKner(ex Natterer), 1860: 4 (published as a synonym of Chalceus orbygnianus; not available).
Megalobrycon erythropterus Cope, 1872: 263, pl.10, fig. 2 ( Typelocality: “Ambyiacu”). Brycon microlepis Perugia, 1897: 149–150 (Type locality “Puerto 14 de Mayo, dipartimento di Bahia Negra, nel Chaco boreale”[Paraguay]); Boulenger, 1898b: 127(same locality); Bertoni, 1914: 11 (Paraguay) (non vidi); Bertoni, 1939: 55(Paraguay); Géry & Mahnert, 1992: 803–806, figs. 6–7 (discussion, short description, literature compilation; Paraguay: “Arroyo Tagatya-mi, est de Puerto Max”; “Concepcion, gué de l’Arroyo Tagatya-mi”; “gué de l’Arroyo Tagatya-guazu”; “Concepcion, Laguna Negra, 15 kmE. de Paso Bareto”; “Concepcion, Estância Primavera”; “en face de Puerto Iguazu”; Brazil: “Lagoa de Sinhá Mariana, 30 kmen aval de Barão de Melgaço”; “Barão de Melgaço, rio Cuiabá”); Silimon et al., 1996: 12(common name); Resende et al., 1998: 9(diet, rio Paraguai basin, Brazil); Margarido & Galetti Jr., 1999: 357– 358 (karyotype); Sabino & Sazima, 1999: 309–312, figs.1–2 (feeding habits, Bonito, Mato Grosso do Sul, Brazil); Britski et al., 2007: 42–43 fig. (description; Brazil, rio Paraguai basin, Pantanal, Brazil); Menezes et al., 2000: 292 (Upper rio Miranda, Mato Grosso do Sul, Brazil); Mateus & Estupiñan, 2002: 165–170 (Rio Cuiabá, Mato Grosso; stock assessment); Wantzen et al., 2002: 242–243, 246–247 (Lake Coqueiro, rio Cuiabá, Mato Grosso; seasonal shifts in stable carbon and nitrogen); Resende, 2003: 119–120, 127, 147 (Rio Paraguai basin; fisheries); Lima, 2003: 176(primary type material listed; as a synonym of Brycon. hilarii); Menni, 2004: 74(listed, Argentina); Mateus et al., 2004: 219(Rio Cuiabá basin, Mato Grosso; fisheries). Brycon orbygnyanus(not Valenciennes): Boulenger, 1897b: 4(Bolivia: “Mission de San Francisco, Rio Pilcomayo; Caiza, Chaco Bolivien”); Ramlow, 1989: 10(Paraguay: Depto. Amambay, Parque Nacional Cerro Corá; Depto. Central, Playa Carrasco). Brycon hilarii: Boulenger, 1900: 3(“Carandasiñho, près de Corumbá”); Eigenmann & Ogle, 1900: 3(Paraguay); Fowler, 1932: 346, 357–358 (Descalvados, Mato Grosso, Brazil); Bertoni, 1939: 55(listed, Paraguay); Miranda-Ribeiro, 1940: 44(Rio Miranda, Salobra, Mato Grosso [do Sul]); Aguirre, 1945: 38, fig. 15 (Rio Paraguai basin, Mato Grosso); Amaral Campos, 1950: 140(part; Mato Grosso); Howes, 1982: 31(Brazil, rio Paraguai, “Pan de ezucar”); Ferraz de Lima, 1987: 89(fisheries, rio Cuiabá, Mato Grosso, Brazil); Sazima & Machado, 1990: 24(predation by piranhas; Pantanal, Poconé, Mato Grosso, Brazil); Barrella et al., 1994: 14–15, 19 (Rio Manso, upper rio Paraguai basin, Mato Grosso, Brazil; ecomorphology, abundance); Beaumord & Petrere Jr., 1994: 26, 32 (same as previous); Agostinho & Júlio Jr., 1999: 379(Paraná, rio Paraná, downstream Itaipu dam, Brazil; common name); Lima, 2003: 176(synonimic list, maximum length, distribution); Makrakis et al., 2007: 191(Canal da Piracema, Itaipu dam rio Paraná); Graça & Pavanelli, 2007: 77(upper rio Paraná, Paraná; short description, picture); Sanches & Galetti, 2007: 889–895 (Rio Miranda, Mato Grosso do Sul: population structure analysed through RAPD markers); Reys et al., 2009: 136–141 (frugivory and seed dispersal, rio Miranda basin, Mato Grosso do Sul); Lima & Ribeiro, 2011: 151, 158 (as an example of “foreland basin” distributional pattern); Bessa et al., 2011: 351–354 (Rio Cuiabá and Rio Miranda basins, upper Paraguai basin, Brazil; aggressive mimicry by Salminus brasiliensis); Resende, 2011: 471, 483–485, 488 (Pantanal, Brazil; reproductive biology); Oliveira et al., 2011: 62–70 (Brazil, Mato Grosso, rio Cuiabá and Manso dam: larval development); Sanches & Galetti, 2012: 408-417 (Rio Miranda, Mato Grosso do Sul: population structure analysed through microsatellite loci). [not La Monte, 1935: 7; Amaral-Campos, 1950: 142; Marlier, 1968: 56; Braga, 1982: 175–180]. Brycon hilarlii(sic): Eigenmann & Kennedy, 1903: 523–524 (Arroyo Trementina, Paraguay). Brycon erythropterus: Fowler, 1939: 263(Contamana, Peru); Eigenmann & Allen, 1942: 254(Rio Ucayali, Orellana; Lago Cashiboya; Rio Pachitea; Iquitos); Ortega & Vari, 1986: 7(listed, Peru; common name); Howes, 1982: 25–26 (literature compilation); Géry & Mahnert, 1984: 176(comparison with B. melanopterus); Géry & Mahnert, 1992: 798, 800, 802–803, fig. 4 (diagnosis, Peru: Marañon, Ucayali; Bolivia: upper rio Madeira); Chang, 1998: 20, 22 (Zona Reservada Tambopata- Candamo, bacia do río Tambopata, Peru). [not Goulding et al., 1988: 124]. Bryconcf. whitei(not Myers & Weitzman): Saul, 1975: 103(Río Aguarico, Napo, Ecuador; diet). Brycon erythropterum: Ortega, 1996: 464(Río Manu, Parque Nacional Manu, depto. Madre de Dios and Cusco, Peru).
Diagnosis. Brycon hilariican be distinguished from all remaining cis-andean Brycon, except for B. orthotaenia, B. orbignyanus, B. whitei, and B. polylepisby possessing a broad caudal peduncle stripe extending into middle caudal-fin rays (vs. caudal peducle and caudal fin color pattern presenting a either a crescent-shaped blotch, an oblique stripe extending into the upper caudal-fin lobe, or no definite color pattern). It can be distinguished from Brycon polylepisby possessing wavy longitudinal stripes along the body, formed by pigmentation concentrated on upper and lower portions of scales (vs. narrow straight longitudinal stripes along the body, formed by pigmentation concentrated in the medio-distal portion of scales), and symphyseal teeth behind the main series of dentary teeth small, considerably smaller than symphyseal teeth belonging to the main series situated immediately in front of it (vs. symphyseal teeth behind the main series of dentary teeth well-developed, approximately as large as symphyseal teeth belonging to the main series situated immediately in front of it). It can additionaly be distinguished from Brycon orbignyanusand B. orthotaeniaprimarily by possessing higher scale counts (67–82, modally 74, vs. 52–63, modally 56, and 49–58, modally 52 lateral line scales in Brycon orbignyanusand B. orthotaenia, respectively; 12–17, modally 15, vs. 10–13, modally 12 and 9–12, modally 10 scales between lateral line and dorsal-fin basis in B. orbignyanusand B. orthotaenia, respectively; 20–28, modally 26, vs. 19–23, modally 20, and 18–21, modally 19 circumpeduncular scales in B. orbignyanusand B. orthotaenia, respectively). Brycon hilariican be distinguished from B. whiteiby presenting a midlateral dark stripe confined to the posterior half portion of body (vs. midlateral dark stripe generally extending anteriorly to supracleithrum area) and by lacking tiny stripes on caudal-fin rays (vs. tiny stripes present in B. whitei). See item “Comparisons” of Brycon amazonicus, for a discussion on the distinction between B. hilariiand the latter species in the Peruvian Amazon.
Description.Morphometric data are presented in Table 16. Large-sized species, largest examined specimen 514.0 mm SL. Body moderately slender in specimens up to 240 mmSL, moderately high in larger specimens. 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 to pronouncedly 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. Seven (2), 8 (12), 9 (12), 10 (17), 11 (16), 12 (5), or 13 (2) relatively small tri- to pentacuspidate teeth in outer series. Four (15), 5 (41), or 6 (11) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 3 (17), 4 (45), or 5 (5) 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 penta- to hexacuspidate, symphyseal teeth tetracuspidate. Maxillary margins approximately parallel, straight in profile. Eleven to 21 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tri- to pentacuspidate, posterior teeth unicuspidate. Dentary with 7 (1), 8 (3), 9 (7), 10 (8), 11 (15), 12 (7), 13 (6), or 14 (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 10–18 small, aciculated, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of fifth to tenth main series dentary teeth. TABLE 16.Morphometric data of Brycon hilarii(A: lectotype, MNHN A.8616). Mean and range does not include lectotype (which is a stuffed specimen). A n Range Mean Standard length (SL) 423.2 68 83.2–514.0 - Percentages of standard length Depth at dorsal-fin origin 25.2 62 25.6–36.1 31.4 Snout to dorsal-fin origin 44.4 68 45.8–53.3 49.7 Dorsal-fin base length 12.5 68 10.1–13.5 11.8 Posterior terminus of dorsal fin to adipose fin 24.8 68 21.2–27.3 24.3 Posterior terminus of dorsal fin to hypural joint 42.0 68 33.6–43.5 37.9 Snout to pelvic-fin insertion 48.6 61 42.7–51.0 46.3 Snout to anal-fin origin 71.3 64 64.6–71.6 67.6 Anal-fin base length 19.5 68 18.9–24.8 22.4 Caudal peduncle length 13.5 68 12.3–18.0 15.5 Dorsal-fin height - 66 16.3–24.6 19.9 Pectoral-fin length - 68 16.7–22.0 19.4 Pelvic-fin length - 60 10.9–22.0 15.6 Caudal peduncle depth 9.1 68 8.5–11.6 10.1 Head length 24.9 68 20.0–29.0 24.6 Percentages of head length Head height 90.8 67 72.0–93.8 83.0 Snout length 31.6 68 27.5–38.2 32.0 Upper jaw length 45.3 68 43.2–52.0 47.7 Horizontal eye diameter 20.4 68 15.5–30.4 23.6 Post-orbital length 48.1 68 43.2–54.5 48.0 Least interorbital width 47.4 68 41.2–53.3 45.9 Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Sixty-seven (2), 68(3), 69(3), 70(3), 71(7), 72(6), 73(8), 74(11), 75(9), 76(4), 77(3), 78(8), 79(2), 80(1), 81(5), or 82 (1) scales in lateral line series. Laterosensory tube simple in specimens smaller than 100 mmSL, ramified in specimens larger than 100 mmSL. Tubules ramification increasing in complexity along ontogeny, specimens up to 150 mmSL with tubules with two or three branches, three to four branches in specimens between 150–230 mmSL, and with more than 10 branches and developing a dendritic pattern of ramification, with tubules overlapping each other in larger (> 300 mmSL) specimens. Horizontal scale rows between dorsal-fin origin and lateral line 12(2), 13(13), 14(15), 15(31), 16(13), or 17(2). Horizontal scale rows between lateral line and pelvic-fin 6(3), 7(18), 8(44), or 9(9). Circumpeduncular scales 20 (1), 21 (5), 22(13), 23(5), 24(4), 25(10), 26(16), 27(12), or 28(9). Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 13th (6) to 14th (2) vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in cs specimens), 19(3), 20(3), 21(7), 22(10), 23(16), 24(14), 25(11), 26(6), 27(2), or 28(2). First anal-fin pterygiophore inserting behind haemal spine of 25th (4) or 26th (4) vertebra. 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 20–26 rectangular scales. Pectoral-fin rays i, 11 (2), 12 (8), 13 (49), 14 (12), or 15 (3). Pelvicfin rays i, 7. Main caudal-fin rays 10/9. Caudal fin slightly forked, distal margin slightly concave. Centralcaudalfin rays presenting in some specimens with a small, pointed middle projection extending beyond primary margin of fin. 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 12 (1), 13 (2), 14 (2), 15 (4), 16 (8), 17 (7), or 18 (2) lower, 1 at angle, and 14 (4), 15 (3), 16 (13), 17 (3), or 18 (3) upper gill rakers. Vertebrae 45 (5), 46 (7), 47 (4), 48 (2), or 49 (1). Supraneurals 10 (6) or 11 (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 fifth to 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. Stripes more discernible dorsally. Series of irregular, narrow vertical stripes present in small (up to 120 mmSL) specimens. Caudal peduncle with broad median stripe, originating 8–12 scales from hypural joint and continuing posteriorly over 6 central principal caudal-fin rays to caudal-fin distal margin. Some specimens with dark stripe extending as a faint stripe anteriorly to the level of anal-fin origin. Remaining caudal-fin rays, and remaining fins, clear, pectoral fins darkened in some specimens. FIGURE 57. Brycon hilarii, lectotype, MNHN A.8616, 423.3 mm SL: Brazil, “Rio San Francisco”. Photo by A.M. Zanata. FIGURE 58. Brycon hilarii, MZUSP 81126, 514.0 mm SL: Brazil, Mato Grosso, rio Cabaçal basin. FIGURE 59. Brycon hilarii, MZUSP 18861, 243.4 mm SL: Brazil, Mato Grosso do Sul, rio Taquari. FIGURE 60. Brycon hilarii, MZUSP 95016, 176.2 mm SL: Brazil, Mato Grosso, rio Mutum. FIGURE 61. Brycon hilarii, MZUSP 26064, 213.6 mm SL: Peru, Ucayali, Río Huacamayo. Coloration in life.Description based on photographs of freshly caught or living specimens from rio Formoso basin, Bonito, Mato Grosso do Sul, Nobres, rio Manso basin, and Nortelândia, rio Santana basin, Mato Grosso, Brazil, from the middle rio Paraná, Corrientes, Argentina, and from Parque Nacional Yanachaga-Chamilen, Río Pachitea drainage, Depto. Pasco, Peru. Overall coloration silvery, dorsum dark-grey, with a coppery hue. Specimens from the rio Paraná and rio Paraguai basins with yellow pigmentation along the lower portion of head and body. Top of head dark grey. Midlateral dark stripe commencing at vertical that passes through origin to middle portion of anal fin, extending through caudal peduncle into middle caudal-fin rays very conspicuous. Caudal-fin rays dorsal and ventral to middle dark stripe reddish to intense red. Anal, dorsal, pectoral, and pelvic fins reddish to intense red. Variation.Specimens from the La Platasystem possess lower total vertebrae counts (45–47, modally 46, n = 14) when compared with specimens from the upper Amazon (47–49, modally 47–48, n = 5). Specimens from the La Platasystem are also on average more high-bodied than specimens from the upper Amazon, as reflected in body depth (27.5–36.1 % SL, mean 32.0, vs. 25.6–33.1 % SL, mean 30.2, respectively) and head height (78.7–93.8 % HL, mean 84.9, vs. 72.0–84.6 % HL, mean 77.7, respectively). Color pattern in life also seems to differ, with specimens from the upper Amazon basin possessing fins with a less intense red pigmentation. Further studies including another typeof data (e.g., molecular data) are necessary to confirm whether both populations are in fact conspecific or not. Sexual dimorphism.None of the examined specimens possess anal-fin hooks. Common names. Brazil: “piraputanga” ( Silimon et al., 1996: 12; Britski et al., 2007: 42); “salmão-criolo” ( Agostinho & Júlio Jr., 1999: 379); Argentina: “salmón de Río”, “pirapitá amarillo” (F. Baena, pers. comm.); Peru: “Sábalo cola roja” ( Ortega & Vari, 1986: 7, as Brycon erythropterum); “Rumi-uma” ( Eigenmann & Allen, 1942: 254, as Brycon erythropterum).
Distribution. Rio Paraguaiand middle rio Paranábasins in Braziland Paraguay, and upper Rio Amazonasbasin in Peru, Ecuador, and Colombia( Fig. 62). Brycon hilariiis recorded for the rio Paranábelow the now submerged Sete Quedas falls, the previous limit between the upper and lower portions of the rio Paraná. Photographs of specimens caught in the rio Paranábetween Paso de la Patria and Itatí, Corrientes, provided by M. Ribeiro and F. Baena, and juveniles in the area of Rosario (province of Santa Fé, c. 32°57’S, 60°40’W), provided by F. Baena, testify the occurrence of Brycon hilariialso in Argentina, from where the species was in fact previously recorded by Menni (2004: 74, as B. microlepis)but not in other check-lists of the ichthyofauna of Argentina( López et al., 2003; Koerber, 2009). We have not examined specimens from Brycon hilariifrom upper tributaries of the Amazon basin in Colombia, but a photograph of a specimen collected at the upper Río Putumayoprovided by A. Linares confirm the occurrence of the species in that country. Although Géry & Mahnert (1992: 800)record Brycon hilariifrom the upper rio Madeirasystem in Bolivia, we have only examined specimens of B. amazonicusfrom this latter area and consider that the eventual occurrence of the species at the upper Rio Madeirabasin in Bolivianeeds further corroboration. The species is being extensively stocked in eastern Braziland accidental/intentional releases in the upper rio Paranábasin are reported ( Graça & Pavanelli, 2007: 77). Ecological notes. Brycon hilariiis a relatively eurytopic species in the rio Paraguaibasin in Braziland Paraguay, occurring from the large rivers in the lowlands and associated floodplains to relatively small, fastflowing rivers in the headwaters. At the middle rio Paranábasin in Braziland Argentina Brycon hilariioccurs syntopically with B. orbignyanus. The largest specimen recorded in the literature measured 670 mmTL ( Mateus & Estupiñan, 2002: 166). Diet and feeding behavior in the rio Paraguaibasin in Brazilwere studied by Sabino & Sazima (1999)and Reys et al.(2009). As other Bryconspecies, B. hilariiis an omnivorous fish that ingests mainly vegetal matter ( Reys et al., 2009). Seeds and fruits constitute an important portion of its diet (31% of the weight), and twelve fruit/seed species were either found in guts or observed to be eaten by Brycon hilariiin the rio Formoso ( Reys et al., 2009). Reys et al.(2009)hypothesized that Brycon hilariiacted as a potential seed disperser for eight tree species that possessed relative small, hard seeds that were not found crushed in the fishes’ guts. Sabino & Sazima (1999: 310–312)observed schools of Brycon hilariiconsisting in up to 30 specimensfollowing capuchin monkeys ( Cebus apella) to catch fruits (specifically, Guareacf. guidonia, Meliaceae, and Zanthoxyllum riedelianum, Rutaceae) that have fallen into the water due to the activity of the monkeys. In the Pantanal area, Resende et al.(1998: 9) reported that Brycon hilarii(their B. microlepis) commonly ingesting flowers of Tabebuia impetiginosa(Bignoniaceae)and pods of Inga uruguensis(Fabaceae). Wantzen et al. (2002: 242, 246–247), noticed an increase in the presence of the isotope 13C in Brycon hilarii(their B. microlepis) during the flooding period, which was correlated to a increased ingestion of terrestrial invertebrates, attesting the importance of allochthnous resources for the species. Considering its importance to fisheries, it is remarkable the lack of studies on the reproduction of the species. As most of its congeners, Brycon hilariiundertake upstream migratory during the rainy season, and juveniles of the species are collected in the flooded areas of the Pantanal between December and March (F. A. Machado, pers. comm.). The population occurring in clear-water upper tributaries of the rio Mirandain Mato Grosso do Sulstate is said to migrate downstream into the turbid, relatively warm rio Mirandato spawn ( Resende, 2011). The species is one of most valued and important targets of both commercial and recreational fisheries of the rio Paraguaibasin in Brazil(Ferraz de Lima, 1987; Resende, 2003; Mateus et al., 2004; Mateus et al., 2011). Analyses of microsatellites loci and RAPD markers in populations of B. hilariifrom the rio Mirandaby Sanches & Galetti (2007, 2012) indicated the presence of a considerable population variability and the presence of specimens distinct from the ones found at local populations in migratory schools, which indicate a considerable population structure, which is unexpected for a migratory species.
Remarks.Valenciennes (in Cuvier & Valenciennes, 1850: 247–248) described Chalceus hilariibased on a specimen collected by Auguste de Saint-Hillaire at the “Rio San Francisco”, and also in “plusieurs autres … dans l’Amazone et dans d’autres rivières du Brésil”, collected by Castelnau. The species was primarily diagnosed from its congeners according to Valenciennes (in Cuvier & Valenciennes, 1850) by its small-sized scales—the specimen from the rio São Francisco possessing 80 lateral-line scales. Bertin (1948: 14–15)identified three syntypes for the species, the specimens MNHN A.8616 (Rio São Francisco; A. St. Hillaire) and MNHN A.9893-9894 (“Amazone”; Castelnau). Géry & Mahnert (1992: 815)selected the specimen MNHN A.8616 as the lectotype of the species, and noticed that the paralectotypes of the species were too poorly preserved to confirm their conspecificity with the lectotype. As noticed under the item “Remarks” of Brycon falcatus, the examination of the typical series of Chalceus hilariishowed that these specimens clearly are not conspecific with the lectotype but rather are referrable to B. falcatus. After its original description, Brycon hilariiwas only reported for the rio Paraguai basin (e.g. Boulenger, 1900; Eigenmann & Ogle, 1907; Fowler, 1932; Miranda-Ribeiro, 1940; Aguirre, 1945; Amaral Campos, 1950). Perugia (1897: 149–150)described Brycon microlepis, from Bahia Negra, Chaco, Paraguay, based on a single specimen (MSNG 36916). This name was considered as valid in the subsequent literature, though virtually ignored until Géry & Mahnert (1992: 806, 814–816)considered it to be the proper name for the Bryconspecies occurring in the rio Paraguai basin. They considered that the name Chalceus hilariicould not possibly refer to the species from the rio Paraguai basin, first because of its stated type-locality (rio São Francisco), and second, because the lateral-line count of the lectotype (60–65) was actually considerably lower than the one mentioned on the original description (80). Géry & Mahnert (1992: 871)suggested instead that Chalceus hilariimight be a senior synonym of Brycon orthotaniafrom the rio São Francisco. This change in name usage, from Brycon hilariito B. microlepis, was followed by authors who subsequently published on the species (e.g. Sabino & Sazima, 1999; Britski et al., 2007; Mateus & Estupiñan, 2002; Resende, 2003; Mateus et al., 2004). However, Lima (2003: 176)reverted to the usage of Brycon hilariifor the species from the rio Paraguai, and considered B. microlepisto be a synonym of it. This change in opinion in spite to the apparently sound remarks by Géry & Mahnert (1992)was prompted by the examination of the lectotype, which actually have higher lateral-line counts than those reported by the aforementioned authors (more than 70). The examination of large series of specimens from the rio São Francisco herein refered as B. orthotaeniashowed a range of 49–58 lateral-line scales, while specimens from the rio Paraguai possess 67–82 lateral-line scales. Also, the overal body shape of the embalmed lectotype of Chalceus hilariiis much more similar to Bryconspecimens from the rio Paraguai basin, than to the species occurring at the rio São Francisco basin. We thus consider that the name Chalceus hilariishould in fact be applied to the Bryconspecies occurring in the rio Paraguai basin, and as such we reject Géry & Mahnert’s (1992) opinion that this name refers to the species from the rio São Francisco basin. However, this leaves us with the problem of the type-locality of Chalceus hilarii, which was stated as being the rio São Francisco by Valenciennes (in Cuvier & Valenciennes, 1850). The botanist Auguste de Saint-Hillaire, the collector of the lectotype of Chalceus hilarii, travelled extensively through eastern and southern Brazil and Uruguay, but never visited the middle rio Paraná or the rio Paraguai ( Papavero, 1971; map 10). The only explanation we can offer to account for this discrepancy is that the lectotype of Chalceus hilariiwas not actually collected by Saint-Hillaire himself, but instead by some other French naturalist that traveled into the rio Paraguai basin at that time (perharps Alcide d’Orbigny), and was at some point mislabeled. It is interesting to note that another fish species described by Valenciennes using material supposedly collected by Saint-Hillaire in the rio São Francisco, Spatuloricaria nudiventris, was never again collected in that river basin, while the genus Spatuloricariais known from the rio Paraguai basin. The systematics of Spatuloricariaare currently still in need of review ( Fichberg et al., 2014), and it is uncertain if the holotype and so far only known specimen of Spatuloricaria nudiventriscorresponds to the Spatuloricariaspecies occurring in the rio Paraguai basin (which is usually identified as S. evansii, a name available for that area). We suggest that both the lectotype of Chalceus hilariiand the holotype of Loricaria nudiventriswere mislabeled in the same event, and that both probably were actually collected somewhere in the rio Paraguai basin. Cope (1872: 262–263)described Megalobrycon erythropterum, as having its type locality at the “Ambyiacu” (= Río Ampyiacu), a small tributary of the Río Amazonas, near Pebas, Loreto, Peru. It is not clear whether Cope had one or more specimens available when he described the species. The type material was not found at the ANSP collection and is presumably lost ( Böhlke, 1984). Megalobrycon erythropterumwas distinguished from the species then assigned by Cope to the genus Megalobrycon( M. cephalusand M. melanopterum) by the “form of body, position of dorsal fin, anal radii, and dentition”. Several authors (e.g., Fowler, 1939; Eigenmann & Allen, 1942; Géry & Mahnert 1984, 1992; Ortega & Vari, 1986) considered it to be a valid species (as Brycon erythropterus). The color plate of the species presented by Cope (1872: pl. 10, fig. 2)and the lateral-line scales counts (estimated by him to be around 70) undoubtedly shows that this name is referrable to the Brycon hilariipopulations from the upper Amazon basin, and, consequently, Megalobrycon erythropterumis herein considered as a synonym of the latter species.
Material examined. Type material:MNHN A.8616 ( 1, 423.3 mmSL): “Rio San Francisco” [type-locality very probably incorrect, see Discussion, above]; A. Saint-Hilaire, no date. Lectotype of Chalceus hilariiValenciennes, designated by Géry & Mahnert (1992: 815). Non types. Brasil, rio Paraguaibasin. Mato Grosso: MZUSP81773( 1, 267.4 mmSL): Nobres, rio Mansobasin, c. 14°45’S, 56°19’W; J.M. Mendes, no date. MZUSP81126(1, 514.0 mm SL): Reserva do Cabaçal, rio do Peixe, trib. rio Cabaçal, 14°55’16’’S, 58°27’59’’W; F.C.T Lima et al., 7 Mar 2002. MZUSP90089( 1, 103.5 mmSL): Cáceres, rio Sepotuba, near its mouth at rio Paraguai, 15°53’34’’S, 57°38’44’’W: H. A. Britski et al., 1 Mar 2002. MZUSP18651(1, 252.0 mm SL): Cáceres, rio Paraguai, 16°4’S, 57°41’W; CEPIPAM, 17–19 Feb 1976. CAS18358 (1, 244.0 mm SL): rio Paraguai, São Luiz de Cáceres, 16°4’S, 57°41’W; J.D. Haseman, 27 May 1909. MCP15817 (2, 144.8– 184.2 mmSL): Cáceres, rio Paraguaiat Cáceresand surroundings, 16°4’S, 57°41’W; R.E.Reis et al., 11 Aug 1991. MCP10752 ( 1, 174.3 mmSL): rio Pixaim, Transpantaneiraroad, 60 kmsouth from Poconé, 16°44’52’’S, 56°51’28’’W; L.R. Malabarba& R.E. Reis, 2 Sept 1986. LISDEBE uncat. (4, 220.0–225.0 mm SL): Poconé, marginal lagoons of Transpantaneiraroad, c. 16°26’S, 56°40’W; J.C. Garavello et al., 16–22 April 1981. ZUEC6853(2, 128.8– 157.7 mmSL): Poconé, Transpantaneiraroad, km 10, c. 16°28’S, 56°41’W; I. Sazima& F. A. Machado, 27 Apr 1981. ZUEC3207( 1, 107.3 mmSL): Poconé, rio Piraputanga, fazenda Ipiranga, c. 16°30’S, 56°45’W; F. A. Machado et al., 16 May 1997. MZUSP19240( 1, 193.9 mmSL): Poconé, c. 16°29’S, 56°41’W; G. Schaller, 1978. MNRJ20525(1, 86.0 mm SL): Chapada dos Guimarães, Água Fria, rio Água Fria, MT-020, 15°10’50’’S, 55°44’52’’W; F. A.G. Melo & M.R.S. Melo, 17 Feb 2000. MZUSP4379( 1, 151.5 mmSL): Santo Antônio do Leverger, rio Cuiabá, 15°52’S, 56°4’W; G. Olson, 1965. MZUSP95016( 1, 176.2 mmSL): Barão de Melgaço, rio Mutum, between Mimoso and Joselândia, 16°19’30’’S, 55°49’59’’W; F. A. Machado et al., 30 Sept 2006. MZUSP89522(2 skel., 272.0–280.0 mm SL): Santo Antônio do Leverger, rio Cuiabá, near mouth of rio Aricá-Açu, Barra do Aricá, 15°59’13’’S, 55°55’34’’W; F. A. Machado, April 2005. ZUEC5705( 1, 130.7 mmSL): Santo Antônio de Leverger, flooded areas near RioCuiabá, c. 15°53’S, 56°2’W; F. A. Machado et al., 28 May 1989. MZUSP85579(8, 3 cs, 71.2–124.3 mmSL): Barão de Melgaço, flooded areas near Mimoso, c. 16°17’S, 55°49’W; F. A. Machado, 19 Jan 1999. MZUSP18795(8, 273.2– 304.4 mmSL): Barão de Melgaço, rio Cuiabá, 16°11’S, 55°58’W; CEPIPAM, 1–10 May 1977. INPA16369( 1, 161.9 mmSL): BaíaSinhá Mariana, c. 30 kmbelow Barão de Melgaço, c. 16°20’S, 55°54’W; CEPIPAM, 30 Nov 1979. MZUSP 67189 (71, 5 cs, 23.4–45.3 mmSL): Barão de Melgaço, flooded areas at Fazenda do Poli; K. de Silimon, 19 Dec 1983. ZUEC5680( 1, 149.9 mmSL): Cuiabá, rio Cuiabá; F. A. Machado, 2 May 1981. FMNH76445(1, 196.0 mm SL): Rio Paraguai, Descalvados, 16°43’S, 57°45’W; Schmidt& Sanborn, 20 Sept 1926. MZUSP27178(2, 192.6– 199.9 mmSL): rio Paraguai, ilha de Taiamã, c. 16°50’S, 57°34’W; A.S. Soares, 1–7 Dec 1980. MZUSP35871( 1, 190.7 mmSL): Itiquira, BaíaGrande, fazenda Santo Antônio do Paraíso, c. 17°33’S, 55°16’W; J.H.B. Medeiros& J.C. Oliveira, 30 April 1979. MZUSP25284( 1, 180.3 mmSL): Itiquira, BaíaGrande, fazenda Santo Antônio do Paraíso, c. 17°33’S, 55°16’W; J.C. Oliveira, 24–29 Oct 1978. Mato Grossodo Sul: MZUSP18681( 1, 243.4 mmSL); MZUSP38206(1, 364.0 mm SL): Coxim, rio Taquari, Cachoeira das Palmeiras, 18°21’45’’S, 56°36’45’’W; CEPIPAM, 6–8 Dec 1976. MZUSP18556( 1, 170.7 mmSL): Coxim, rio Taquari, c. 18°29’S, 54°45’W; A. Storti & W. Uieda, 15 Aug 1975. MZUSP2910(5, 231.7– 268.1 mmSL): Coxim, rio Piquerí, c. 17°55’S, 54°41’W; J. Lima, Oct 1930. MZUSP17299( 1, 215.1 mmSL): rio Taquari, c. 150 kmfrom Coxim; J.C. Garavello et al., 9–22 Aug 1967. BMNH1910.5.26.19–20 (2, 244.0–284.0 mm SL): “ Pan de Ezucar, R. Paraguay”(not located); C. Grant, no date. MZUSP48297(2, 1 cs, 154.3–161.2 mmSL): Fazenda Santo Antônio( Baíada Sede), Pantanal de Paiaguás, c. 17°53’S, 57°8’W; T. Liparelli, no date. MZUSP 36444 ( 1, 171.9 mmSL): Corumbá, Corixão, Capão Grande, Nhecolândia; G.M. Mourão et al., Sept 1985. MZUSP18936( 1, 247.8 mmSL): rio Paraguai, surroundings of Corumbá, 19°00’S, 57°38’W; CEPIPAM, 12 Oct 1977. CAS18359 ( 1, 257.8 mmSL), lagoons and rivers near Corumbá, 19°0’S, 57°38’W; J.D. Haseman, 28 April 1909. MZUSP63173( 1, 221.8 mmSL): Corumbá, rio Paraguai, Baíade Albuquerque, 19°24’54’’S, 57°22’43’’W; CEPIPAM, 19 Feb 1977. MZUSP49023(3, 13.8–18.4 mmSL): Corumbá, Passo do Lontra, 19°34’S, 57°1’W; Exp. Zool. UFMS, 13 Jan 1988. LBP 37 (3, 141.3– 151.9 mmSL): Corumbá, rio Miranda, Passo do Lontra, 19°34’38’’S, 57°01’8’’W; C. Oliveira et al., July–Aug 1996. MZUSP 59529 (1, 166.0 mm SL); MZUSP59527( 1, 181.6 mmSL): Corumbá, rio Abobral2, 19°26’68’’S, 57°02’49’’W; A. Machado-Allison et al., 7 Sept 1998. MZUSP59528( 1, 133.2 mmSL): Aquidauana, fazenda São Pedro, 19°36’70’’S, 56°02’78’’W; A. Machado-Allison et al., 4 Sept 1998. MZUSP2986( 1, 232.3 mmSL): Miranda, Salobra, rio Miranda, 20°12’S, 56°30’W; L. Travassos, Aug 1940. MNRJ2878(1, 286.0 mm SL): Miranda, rio Miranda, Salobra, 20°12’S, 56°30’W; L. Travassos, 1941. MZUSP3073( 1, 183.7 mmSL): “ rio Bodoquena” (precise locality uncertain, see under Remarksof Brycon orbignyanus); L. Travassos, 1941. Paraná, rio Paranábasin: MZUSP21064(7, 277.4– 357.9 mmSL): Guaíra, rio Paranábelow Sete Quedaswaterfalls, c. 24°7’S, 54°20’W; CETESB, 1977–1978. MZUSP 14652 (1, 374.0 mm SL); MZUSP 14653 (1, 327.0 mm SL); MZUSP14654(1, 330.0 mm SL): PortoMendes, rio Paraná, 24°29’S, 54°19’W; CETESB, 1978. Paraguay, rio Paraguaibasin: FMNH108173(1, 262.0 mm SL): Depto. Alto Paraguay, RioNegro, ca. 2.5 kmabove mouth of the Rio Paraguay, 20°9'S, 58°10'W; D. Mandelburger et al., 5 Sept 1997. FMNH108174(2, 313.0–318.0 mm SL): Depto. Alto Paraguay, Rio Paraguay, above Estancia Cerrito, 21°27'S, 57°55'W; M.T Piza et al., 10 Sept 1997. USNM232317( 1, 257.1 mmSL): Depto. Amambay, Parque Nacional Cerro Cora, RioAquidaban, c. 22°38’S, 56°1’W; L. Naylor& C. Cuevas, 16 Jan 1982. UMMZ206816( 1, 233.8 mmSL): Depto. Concepción, ArroyoCagata, c. 14 kmS of Yby-Yaú, 23°6’30’’S, 56°31’12’’W; J. Taylor et al., 28 Jul 1979. CAS77377 (1, 174.0 mm SL): Depto. Concepción, ArroyoTrementina, trib. of the R. Aquido Canigi(= RioAquidaban?), c. 23°3’S, 57°0’W; J.D. Anisits, Dec 1900. USNM181763( 1, 247.7 mmSL): Depto. Central, Asuncionbay, Río Paraguaynear Asuncion, c. 25°16’S, 57°38’W; C.J.D. Brown, 15 Jan 1957. UMMZ208068( 1, 278.8 mmSL): Depto. Central, Río Paraguay, overflow c. 1 kmbelow Puerto Remanzobridge, c. 25°11’S, 57°33’W; J.Taylor et al., 15 Sept 1979. USNM181762(2, 236.2–239.0 mm SL): Depto. Cordillera, ArroyoPirareta, near Piribebuy, c. 25°30’S, 56°56’W; C.J.D. Brown, 13 Dec 1956. UMMZ205665( 1, 237.1 mmSL): Depto. Cordillera, Saltode Pirareta, c. 400 mbelow fall, c. 25°30’S, 56°56’W; J.N. Taylor et al., 25–26 May 1979. USNM181761( 1, 234.3 mmSL): RíoTebicuary near “ Reciefe” (not located); C.J.D. Brown, 6 Dec 1956.
Rio Amazonasbasin, Peru. Depto. Loreto: MZUSP 26507( 1, 228.7 mmSL): RíoUcayali, “ Albufao” (not located); J. Guevara, 9 Oct 1971. NRM23675 ( 1, 230.3 mmSL): El Estrecho, Río Putumayo, 2°27’S, 72°40’W; S.O. Kullander et al., 18 Jul 1986. MUSM12745( 1, 132.6 mmSL): Yanayacu, Base3, Pluripetrol, RíoShiriyacu, 4°51’S, 74°56’W; H. Ortega, 10 Oct 1997. Depto. Ucayali: MZUSP26064( 1, 213.6 mmSL); MUSM 31 (190.0 mm SL); MUSM30(1, 215.0 mm SL): RíoHuacamayo, road Pucallpa-Huánuco(= carretera Aguaytia/ Pucallpa), km 7.5; J. Guevara, 17 Oct 1971. MUSM uncat. ( 1, 190.9 mmSL): Pucallpa, RíoNeshuya, IVITA, 8°38’S, 74°58’W; H. Ortega, 6 Dec 1971. MZUSP26471(1, 83.2 mmSL): RíoNeshuya, road Pucallpa-Huánuco, 8°38’S, 74°58’W; H. Ortega, 23 Jan 1976. MZUSP26373( 1, 108.8 mmSL): RíoSan Alejandro, Pucallpa, c. 8°50’S, 75°13’W; J. Guevara, 14 Nov 1971. MUSM28(1, 248.0 mm SL): RíoSaballo, trib. RíoAguaytia, Aguaytia, Tangarana; J. Guevara, 14 Oct 1971. MUSM28573( 1, 167.4 mmSL): Padre Abad, RíoAguaytia basin, Río Santa Ana, Quebrada Samíria, 8°32’41’’S, 75°38’35’’W; M. Hidalgo& M. Rojas, 5 Dec 2006. MUSM31(2, 248.0– 288.1 mmSL): Portoof Pucallpa, RíoUcayali; H. Ortega, 10 Oct 1971. MUSM381(1, 159,0 mm SL): Pucallpa, Tachsitea, Río Calleria(trib. RíoUcayali), 8°1’S, 74°35’W; P. de Rham& H. Ortega, 4 Oct 1984. MUSM23(1, 165.0 mm SL): Pucallpa, RíoUcayali, Masisea, 8°35’S, 74°20’W; H. Ortega, 24 Sept 1975. MUSM41413(1, 79.1 mmSL): Pucallpa, RíoUcayali, Masisea, 8°35’S, 74°20’W; H. Ortega, 23 Nov 1973. MUSM 3562 (2, 177.3– 219.0 mm SL): Pucallpa, IVITA, Campo Verde; J. Guevara, 11 Oct 1971. FMNH84312(1, 267.0 mm SL): mouth of RioSan Alejandro at junction with Sungaro Yacu, c. 8°27’S, 75°11’W; G. Gutierrez, 1 Aug 1975. FMNH84095(1, 393.0 mm SL): RioSan Alejandro, deep pool upstream from camp, c. 8°27’S, 75°11’W; G.S. Glodek, 3 Aug 1975. FMNH84094(1, 383.0 mm SL): RioSan Alejandro, c. 8°27’S, 75°11’W; D.W. Greenfield, 3 Aug 1975. CAS 16025 ( 1, 275.1 mmSL); CAS68879 (ex IU 16025) (2, 162.7– 166.9 mmSL): Lago Cashiboya, a cutoff lake of RíoUcayali(connected to river by a channel) above Contamana, 7°40’S, 74°56’W; W.R. Allen, 3–4 Aug 1920. CAS68878 (ex IU 16023) (3, 151.8–186.0 mm SL): RíoUcayaliat Orellana, 6°55’S, 75°9’W; W.R. Allen, 9 Aug 1920. Depto. Huánuco: NRM23669 ( 1, 150.2 mmSL): near mouth of quebrada trib. RíoPachitea, right bank, 2– 3 kmdownstream Tournavista, c. 8°57’S, 74°41’W; S.O. Kullander& A. Hogeborn, 19 Aug 1981. CAS68881 (ex IU 16026) ( 1, 196.2 mmSL): RíoPachitea; W.R. Allen, 20–22 Aug 1920. ROM55913 ( 1, 167.3 mmSL): RíoLlullapichis (trib. RíoPachitea), Panguana, RíoPachitea drainage, c. 9°37'S, 74°56'W; H. Pandura, Jul 1988. Ecuador: FMNH103396( 1, 322.2 mmSL): Napo, Tiuyacu, first tributary to Rio Churuayacuupstream from mouth at Rio Payaminoand near mouth of Rio Churuyacu, 0°29'30''S, 77°18'W; D. Stewart, M. Ibarra& R. Barriga, 13 Nov. 1983. MCZ60933 (1, 298.0 mm SL): Napo, RíoPunino, trib. RíoPayamino, above Coca, c. 0°30’S, 77°0’W; T.R. Roberts & H. Pauker, 25 Nov 1971. FMNHuncat. (2, 416.0–426.0 mm SL): Morona- Santiago, Rio Pastaza, brazo lateral at Puerto Pakitza, 2°15’45’’S, 77°14’17’’W; B. Chernoff& R. Barriga, 17–18 Jul 1999.
1425221977
MZUSP
Brazil
Non
-14.75
Nobres
1287
-56.316666
rio Paraguai
105
106
MZUSP 81773
1
Mato Grosso
holotype
1425221972
2002-03-07
MZUSP
J. M. Mendes
Brazil
Reserva do Cabacal
-14.921111
rio Cabacal
21
-58.46639
Peixe
105
106
MZUSP 81126
1
Lima
holotype
1425221975
2002-03-01
MZUSP
Paraguay
-15.892777
rio Sepotuba
21
-57.645557
Caceres
105
106
MZUSP 90089
1
A
holotype
1425221983
[223,1432,583,608]
1976-02-17
1976-02-19
1976-02-17
MZUSP
Paraguay
-16.066668
Caceres
1283
-57.683334
105
106
MZUSP 18651
1
holotype
1425221974
[151,1432,620,645]
1909-05-27
CAS
Sao Luiz de Caceres & J. D. Haseman
Paraguay
-16.066668
Sao Luiz de Caceres
1283
-57.683334
105
106
1
holotype
1425221981
[151,1431,655,680]
MCP
Paraguay
-16.066668
Caceres
1283
-57.683334
Caceres
105
106
1
holotype
1425221976
1991-08-11
MCP
R. E. Reis
Paraguay
rio Pixaim
-16.747778
Pocone
21
-56.857777
Transpantaneira
105
106
1
holotype
1425221995
1986-09-02
L. R. Malabarba & R. E. Reis
Paraguay
-16.433332
Transpantaneira
1282
-56.666668
Pocone
105
106
1
holotype
1425221979
1981-04-16
1981-04-22
1981-04-16
ZUEC
J. C. Garavello
Paraguay
-16.466667
Transpantaneira
1282
-56.683334
Pocone
105
106
ZUEC 6853
1
holotype
1425221989
1981-04-27
ZUEC
I. Sazima
Paraguay
Pocone
-16.5
Ipiranga
1282
-56.75
rio Piraputanga
105
106
ZUEC 3207
1
A
holotype
1425221973
1997-05-16
MZUSP
Paraguay
-16.483334
Pocone
1282
-56.683334
105
106
MZUSP 19240
1
A
holotype
1425221986
MNRJ
G. Schaller & Chapada dos Guimaraes
Paraguay
-15.180556
rio Agua Fria
21
-55.74778
Agua Fria
105
106
MNRJ 20525
1
A
holotype
1425221982
2000-02-17
MZUSP
M. R. S. Melo
Paraguay
-15.866667
rio Cuiaba
1284
-56.066666
Santo Antonio do Leverger
105
106
MZUSP 4379
1
A
holotype
1425221994
MZUSP
G. Olson & Barao de Melgaco
Paraguay
-16.325
rio Mutum
21
-55.833054
Barao de Melgaco
105
106
MZUSP 95016
1
A
holotype
1425221978
2006-09-30
MZUSP
Paraguay
Santo Antonio do Leverger
-15.986945
rio Arica-Acu
21
-55.926113
rio Cuiaba
105
106
MZUSP 89522
1
A
holotype
1425221996
[822,1125,1087,1112]
2005-04
Paraguay
A
105
106
1
A
holotype
1425221988
ZUEC
Santo Antonio de Leverger
Paraguay
-15.883333
Santo Antonio de Leverger
1284
-56.033333
105
106
ZUEC 5705
1
Rio
holotype
1425221991
1989-05-28
MZUSP
Barao de Melgaco
Paraguay
-16.283333
Mimoso
1282
-55.816666
Barao de Melgaco
105
106
MZUSP 85579
1
A
holotype
1425221987
1999-01-19
MZUSP
Barao de Melgaco
Paraguay
-16.183332
rio Cuiaba
1282
-55.966667
Barao de Melgaco
105
106
MZUSP 18795
1
A
holotype
1425221984
1977-05-01
1977-05-10
1977-05-01
INPA
Barao de Melgaco
Paraguay
-16.333334
Barao de Melgaco
1282
-55.9
105
106
INPA 16369
1
Baia
holotype
1425221993
ZUEC
Paraguay
rio Cuiaba
Cuiaba
105
106
ZUEC 5680
1
Baia
holotype
1425221990
[453,1119,1340,1365]
1981-05-02
FMNH
Paraguay
A
105
106
FMNH 76445
1
A
holotype
1425221980
1926-09-20
Schmidt & Sanborn
Paraguay
-16.716667
Descalvados
1281
-57.75
105
106
1
Rio
holotype
1425221985
MZUSP
Paraguay
-16.833334
ilha de Taiama
1280
-57.566666
105
106
MZUSP 27178
1
holotype
1425221992
[544,1371,1412,1437]
1980-12-01
1980-12-07
1980-12-01
MZUSP
Paraguay
A
105
106
MZUSP 35871
1
A
holotype
1425221997
1979-04-30
J. H. B. Medeiros & J. C. Oliveira
Paraguay
-17.55
Santo Antonio do Paraiso
1278
-55.266666
105
106
1
Baia
holotype
1425222018
1978-10-24
1978-10-29
1978-10-24
MZUSP
J. C. Oliveira
Paraguay
-17.55
Santo Antonio do Paraiso
1278
-55.266666
Itiquira
105
106
MZUSP 25284
1
Baia
holotype
1425222001
[505,1143,1519,1544]
MZUSP
Paraguay
Mato Grosso do Sul
105
106
MZUSP 18681
1
Mato Grosso do Sul
holotype
1425222011
1976-12-06
1976-12-08
1976-12-06
MZUSP
Cachoeira das Palmeiras
Paraguay
-18.362501
rio Taquari
21
-56.6125
Coxim
105
106
MZUSP 38206
1
Mato Grosso do Sul
holotype
1425222019
[151,1015,1591,1616]
MZUSP
Paraguay
-18.483334
rio Taquari
1275
-54.75
Coxim
105
106
MZUSP 18556
1
Mato Grosso do Sul
holotype
1425222020
1975-08-15
MZUSP
W. Uieda
Paraguay
-17.916666
rio Piqueri
1277
-54.683334
Coxim
105
106
MZUSP 2910
1
A
holotype
1425221998
[1115,1330,1628,1653]
1930-10
Paraguay
Lima
105
106
1
Lima
holotype
1425222021
1967-08-09
1967-08-22
1967-08-09
MZUSP
Coxim & J. C. Garavello
Paraguay
rio Taquari
105
106
MZUSP 17299
1
Lima
holotype
1425222000
BMNH
Pan de Ezucar, R. & C. Grant
Paraguay
Paraguay
105
106
1
holotype
1425222003
MZUSP
Pantanal de Paiaguas
Paraguay
-17.883333
Pantanal de Paiaguas
1277
-57.133335
Fazenda Santo Antonio
105
106
MZUSP 48297
1
Baia
holotype
1425221999
1977-10-12
MZUSP
Paraguay
-19.0
Corumba
1273
-57.633335
105
106
MZUSP 18936
1
holotype
1425222007
1909-04-28
CAS
J. D. Haseman
Paraguay
-19.0
Corumba
1273
-57.633335
105
106
1
holotype
1425222004
MZUSP
Paraguay
-19.414999
Corumba
21
-57.37861
105
106
MZUSP 63173
1
Baia
holotype
1425222014
1977-02-19
MZUSP
Paraguay
-19.566668
Passo do Lontra
1271
-57.016666
Corumba
105
106
MZUSP 49023
1
Baia
holotype
1425222015
1988-01-13
UFMS
Paraguay
-19.577223
Passo do Lontra
21
-57.01889
Corumba
105
106
1
Miranda
holotype
1425222005
[447,1432,151,176]
MZUSP
Paraguay
-19.45222
rio Abobral
21
-57.046944
Corumba
106
107
MZUSP 59527
1
Miranda
holotype
1425222010
1998-09-07
MZUSP
Paraguay
-19.619444
Sao Pedro
21
-56.055
Aquidauana
106
107
MZUSP 59528
1
A
holotype
1425222009
[468,927,223,248]
1998-09-04
Paraguay
A
106
107
1
A
holotype
1425222013
MZUSP
Paraguay
-20.2
Salobra
1268
-56.5
106
107
MZUSP 2986
1
Miranda
holotype
1425222016
1940-08
MNRJ
L. Travassos
Paraguay
-20.2
Salobra
1268
-56.5
106
107
MNRJ 2878
1
Miranda
holotype
1425222008
MZUSP
L. Travassos
Paraguay
Remarks
rio Bodoquena
106
107
MZUSP 3073
1
Miranda
holotype
1425222012
MZUSP
Paraguay
Parana
106
107
MZUSP 21064
1
Parana
holotype
1425222017
1977
1978
1977
CETESB
Paraguay
-24.116667
Sete Quedas
1252
-54.333332
rio Parana
106
107
1
Guaira
holotype
1425222006
MZUSP
Paraguay
-24.483334
Porto
1251
-54.316666
106
107
MZUSP 14654
1
Porto
holotype
1425222002
FMNH
Paraguay
Depto
rio Paraguai
106
107
FMNH 108173
1
holotype
1425222023
[722,886,476,501]
Paraguay
Alto Paraguay
106
107
1
Alto Paraguay
holotype
1425222041
1997-09-05
FMNH
D. Mandelburger
Paraguay
-20.15
Rio Negro
1268
-58.166668
106
107
FMNH 108174
1
Rio Negro
holotype
1425222035
Paraguay
Depto. Alto Paraguay
106
107
1
Depto. Alto Paraguay
holotype
1425222029
1997-09-10
USNM
M. T Piza
Paraguay
-21.45
Estancia Cerrito
1264
-57.916668
106
107
USNM 232317
1
Rio Paraguay
holotype
1425222030
[536,984,583,608]
Paraguay
Amambay
106
107
1
Amambay
holotype
1425222036
1982-01-16
UMMZ
L. Naylor & C. Cuevas
Paraguay
-22.633333
Rio
1259
-56.016666
106
107
UMMZ 206816
1
Rio
holotype
1425222022
[1048,1185,620,645]
Paraguay
Concepcion
106
107
1
Concepcion
holotype
1425222040
1979-07-28
CAS
J. Taylor
Paraguay
-23.108334
Yby-Yau
21
-56.52
106
107
1
Arroyo
holotype
1425222026
[151,288,692,717]
Paraguay
Concepcion
106
107
1
Concepcion
holotype
1425222039
1900-12
USNM
R. Aquido Canigi & J. D. Anisits
Paraguay
-23.05
Arroyo
1257
-57.0
106
107
USNM 181763
1
Arroyo
holotype
1425222024
[844,929,727,752]
Paraguay
Central
106
107
1
Central
holotype
1425222046
[940,1096,727,752]
Paraguay
Asuncion
106
107
1
Asuncion
holotype
1425222037
1957-01-15
UMMZ
C. J. D. Brown
Paraguay
-25.266666
Rio
1247
-57.633335
106
107
UMMZ 208068
1
Rio
holotype
1425222045
[1293,1378,764,789]
Paraguay
Central
106
107
1
Central
holotype
1425222044
1979-09-15
USNM
J. Taylor
Paraguay
-25.183332
Puerto Remanzo
1247
-57.55
106
107
USNM 181762
1
Rio Paraguay
holotype
1425222043
[703,820,836,861]
Paraguay
Cordillera
106
107
1
Cordillera
holotype
1425222042
1956-12-13
UMMZ
C. J. D. Brown
Paraguay
-25.5
Piribebuy
1246
-56.933334
106
107
UMMZ 205665
1
Arroyo
holotype
1425222028
[994,1111,871,896]
Paraguay
Cordillera
106
107
1
Cordillera
holotype
1425222034
1979-05-25
1979-05-26
1979-05-25
J. N. Taylor
Paraguay
400
-25.5
Salto
1246
-56.933334
106
107
1
Salto
holotype
1425222031
1956-12-06
USNM
Reciefe & C. J. D. Brown
Paraguay
Rio
106
107
USNM 181761
1
Rio
holotype
1425222032
[199,606,980,1005]
Peru
Amazonas
106
107
1
Amazonas
1425222033
[620,709,980,1005]
MZUSP
Peru
Loreto
106
107
MZUSP 26507
1
Loreto
1425222025
1971-10-09
NRM
Albufao & J. Guevara & El Estrecho
Peru
Rio Ucayali
106
107
1
Rio Ucayali
1425222038
1986-07-18
S. O. Kullander
Peru
-2.45
Rio Putumayo
1307
-72.666664
106
107
1
Rio Putumayo
1425222027
1997-10-10
MUSM
H. Ortega
Peru
Yanayacu
-4.85
Pluripetrol
1306
-74.933334
Base
106
107
MUSM 12745
1
Rio
1425222048
[643,1222,1087,1112]
MZUSP
Peru
Depto. Ucayali
106
107
MZUSP 26064
1
Depto. Ucayali
1425222063
1971-10-17
MUSM
Pucallpa-Huanuco & Aguaytia & Pucallpa & J. Guevara
Peru
Rio
106
107
MUSM 30
1
Rio
1425222059
1971-12-06
IVITA
H. Ortega
Peru
-8.633333
Rio
1301
-74.96667
106
107
1
Rio
1425222053
1976-01-23
MZUSP
H. Ortega
Peru
-8.633333
Pucallpa-Huanuco
1301
-74.96667
106
107
MZUSP 26471
1
Rio
1425222058
1971-11-14
MZUSP
J. Guevara
Peru
-8.833333
Pucallpa
1300
-75.21667
106
107
MZUSP 26373
1
Rio
1425222057
[583,1142,1268,1293]
MUSM
Peru
Rio
106
107
MUSM 28
1
Rio
1425222047
1971-10-14
MUSM
Aguaytia & Tangarana & J. Guevara
Peru
Rio
106
107
MUSM 28573
1
Rio
1425222065
[1096,1315,1303,1328]
Peru
Rio
106
107
1
Rio
1425222050
Peru
-8.544723
Quebrada Samiria
21
-75.64305
106
107
1
Rio Santa Ana
1425222064
2006-12-05
MUSM
M. Rojas
Peru
Hidalgo
106
107
MUSM 31
1
Hidalgo
1425222049
[330,529,1375,1400]
Peru
Porto
106
107
1
Porto
1425222067
1971-10-10
MUSM
H. Ortega
Peru
Rio Ucayali
106
107
MUSM 381
1
Rio Ucayali
1425222061
[271,1263,1412,1437]
1984-10-04
P. de Rham & H. Ortega
Peru
-8.016666
Rio Calleria
1302
-74.583336
106
107
1
Rio Ucayali
1425222071
1975-09-24
MUSM
H. Ortega
Peru
-8.583333
Masisea
1301
-74.333336
Pucallpa
106
107
MUSM 23
1
Rio Ucayali
1425222069
1973-11-23
MUSM
H. Ortega
Peru
-8.583333
Masisea
1301
-74.333336
Pucallpa
106
107
MUSM 41413
1
Rio Ucayali
1425222070
1975-08-01
FMNH
G. Gutierrez
Peru
-8.45
Sungaro Yacu
1301
-75.183334
106
107
FMNH 84312
1
Rio
1425222068
1975-08-03
FMNH
G. S. Glodek
Peru
-8.45
Rio
1301
-75.183334
106
107
FMNH 84095
1
Rio
1425222062
[223,1432,1628,1653]
1975-08-03
FMNH
D. W. Greenfield
Peru
-8.45
Rio
1301
-75.183334
106
107
FMNH 84094
1
Rio
1425222060
1920-08-03
1920-08-04
1920-08-03
CAS
W. R. Allen
Peru
-7.6666665
Contamana
1303
-74.933334
Lago Cashiboya
106
107
1
Rio Ucayali
1425222056
1920-08-09
CAS
W. R. Allen
Peru
-6.9166665
Orellana
1303
-75.15
106
107
1
Rio Ucayali
1425222054
[221,1112,1772,1797]
NRM
Peru
Huanuco
106
107
1
Huanuco
1425222055
1981-08-19
S. O. Kullander
Peru
-8.95
Tournavista
1300
-74.683334
106
107
1
Rio
1425222051
1920-08-20
1920-08-22
1920-08-20
CAS
W. R. Allen
Peru
Rio
106
107
1
Rio
1425222066
ROM
Peru
Rio
106
107
1
Rio
1425222052
[665,1432,1879,1904]
1988-07
H. Pandura
Peru
-9.616667
Rio
1299
-74.933334
106
107
1
Rio
1425222073
[151,675,1916,1941]
FMNH
Ecuador
Ecuador
106
107
FMNH 103396
1
1425222087
1983-11-13
MCZ
D. Stewart & M. Ibarra & R. Barriga
Ecuador
Tiuyacu
-0.49166664
Rio Payamino
925
-77.3
Rio Churuayacu
106
107
1
Napo
1425222088
[805,868,1989,2012]
Ecuador
Napo
106
107
1
Napo
1425222080
[882,1072,1988,2013]
Ecuador
Rio
106
107
1
Rio
1425222079
[1085,1436,1988,2013]
Ecuador
Rio
106
107
1
Rio
1425222083
1999-07-17
1999-07-18
1999-07-17
FMNH
Puerto Pakitza & B. Chernoff & R. Barriga
-2.2625
Santiago, Rio Pastaza
21
-77.23805
107
108
1
Santiago, Rio Pastaza