taxonID	type	description	language	source
C033D7104F71FFF64EA4F9C4FB4AFA7A.taxon	diagnosis	Diagnosis. The genus Brycon can be diagnosed from all remaining characid genera, with the exception of Triportheus and Chilobrycon, by the possession of three rows of premaxillary teeth with a distinctive arrangement (see “ Monophyly of Brycon ”, below, for more details). Brycon can be diagnosed from Triportheus for lacking the ventral keel formed by greatly developed coracoid bones extending from the isthmus to a little ahead of the anal-fin origin, characteristic of the latter genus, as well as possessing moderately-sized pectoral fins, reaching only the pelvic-fin origin (vs. pectoral fins elongated and pointed, their tips always beyond pelvic-fin origin in Triportheus) and pelvic-fin originating approximately at the same level than dorsal-fin (vs. pelvic fins situated ahead of dorsalfin origin). Brycon can be distinguished from the closely related and monotypic genera Chilobrycon and Henochilus by possessing broad, bulky dentary and premaxillary teeth (vs. compressed, incisiform teeth; see Géry & de Rham, 1981; Castro et al., 2004), and from Henochilus by possessing three rows of premaxillary teeth (vs. two rows of premaxillary teeth in Henochilus; see Castro et al., 2004). Brycon can be diagnosed from the somewhat similar-looking South American chalceid genus Chalceus by possessing relatively small scales, always more than 40 scales in lateral-line counts, and approximately of the same size across the body (vs. large scales, never more than 40 scales in lateral-line counts, and scales above lateral line considerably larger than those at lateral line or below it in Chalceus), and a relatively long anal fin, with always more than 15 branched rays (vs. short anal fin, with 8 – 10, generally 9, branched anal-fin rays in Chalceus). Additional remarks on the diagnosis and putative monophyly of Brycon are presented below. Monophyly of Brycon. The monophyly of Brycon has been questioned for some time in the literature (e. g., Howes, 1982; Castro & Vari, 1990), and molecular evidence pointing its non-monophyly has recently emerged (Abe et al., 2014; see “ Intrarrelationships ”, below). So far, no unique derived features were identified for the genus Brycon, and consequently, the monophyly of the genus still remains uncertain. Although a resolution on the question of the monophyly of the genus lies beyond the scope of the present contribution, comments on the distribution and variation of three relatively uncommon character-states that are usually employed for diagnosing the genus are presented below, with a view of helping future discussions on its putative monophyletic status.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F71FFF64EA4F9C4FB4AFA7A.taxon	description	1. Three series of premaxillary teeth. Brycon species are unusual among characiforms in typically possessing three series of premaxillary teeth (four series of premaxillary teeth are present in Brycon dentex and some other trans-andean / middle American Brycon species; see Howes, 1982: 21, 23). An interpretation for the teeth arrangement in Brycon, first advanced by Böhlke (1958: 68), employed by Howes (1982: 2), and adopted herein, is to consider the outer series formed by small teeth as the first series, the pair of enlarged teeth near the symphysis of the premaxillary as the third series, and the middle-sized teeth situated at the inner (lingual), lateral border of the premaxillary and between the first, outer series and the inner, third series in the middle portion of the premaxillary as being the second series (Figs. 1 – 2). Three premaxillary teeth rows are also found, among characids, in Bryconops alburnoides (e. g., Machado-Allison et al., 1993), Piabina, Creagrutus (Vari & Harold, 2001: 12 – 13), Triportheus, and Chilobrycon, in the chalceid genus Chalceus and in the alestid Bryconaethiops (Zanata & Vari, 2005: 32 – 33). The premaxillary teeth arrangement found in Creagrutus and Piabina, discussed in detail by Vari & Harold (2001: 12 – 13) and Ribeiro et al. (2004: 609), is very distinct and certainly not homologous to the one found in Brycon. We concur with Zanata & Toledo-Piza (2004: 105) and Zanata & Vari (2005: 32 – 33) in considering the condition found in Chalceus as non-homologous with the one found in Brycon. In Chalceus, the inner series, third series is formed by 5 – 10 teeth that extend from the premaxillary symphysis to the inner, lateral portions of the bone, while the second series is formed by two teeth situated between the first and third teeth rows (Zanata & Toledo-Piza, 2004: 10). This is clearly a distinct condition from the one found in Brycon, where the pair of enlarged teeth does not seem to be closely associated with the smaller teeth situated in the inner, lateral portion of the premaxillary, which are instead distinctly aligned with the similar-sized teeth situated in the middle of the premaxillary, between the outer (first) and inner (third) premaxillary teeth rows (Figs. 1 – 2; see also Howes, 1982: figs. 1, 2, 16, and 18). We have not examined any Bryconaethiops specimens, but the condition in this latter genus, as indicated by Zanata & Vari (2005: 33, 35, fig. 15; see also Roberts & Stewart, 1976: 274, fig. 3, and Teugels & Thys van den Audenaerde, 1990: 210, fig. 4) clearly does not correspond to the one found in Brycon and should also be considered as non-homologous. As for the condition presented by Triportheus and Chilobrycon, we concur with Zanata & Toledo-Piza (2004: 105) and Zanata & Vari (2005: 33) in considering it to be virtually undistinguishable from the condition found in Brycon (see Fig. 3). Mirande (2010: 482) recovered the presence of three series of teeth on the premaxillary as one of the synapomorphies for Bryconinae (in his definition, including, besides Brycon, the genus Triportheus), although when coding this character he disregarded the hypothesis of nonhomology between the condition found in these genera, on one hand, and the one found in Chalceus and Bryconaethiops, raised by Zanata & Toledo-Piza (2004) and Zanata & Vari (2005), which is also supported here. Another synapomorphy for Bryconinae involving the arrangement of the premaxillary teeth proposed by Mirande (2010: 482) was the “ polymorphism of teeth on the inner premaxillary row … with two medial teeth somewhat larger and usually separated from remaining ones by a gap ”. This corresponds to the presence of the inner, third series of teeth in Brycon, which Mirande (2010: 423) considered as also including the teeth present in the inner, lateral portions of the premaxillary. Interestingly, Mirande (2010: 471, 482) suggests that this character state is also present in Serrasalmidae, though the latter only present two teeth rows (with the exception of the piranhas, which possess a single row of premaxillary teeth; Machado-Allison, 1985: 22 – 25; Cione et al., 2009). Premaxillary teeth arrangement in basal Serrasalmidae is in fact remarkably similar to the one observed in Bryconinae not only in the possession of a pair of inner enlarged teeth near the premaxillary series (which corresponds, respectively, to what is generally considered to be part of the second series in Serrasalmidae, and the third series in Bryconinae), but also because the outer, first teeth series of Serrasalmidae has a very similar arrangement to the second teeth series in Bryconinae (compare Figs. 1 – 2 with Fig. 4; see also Jégu et al., 2003: 841, fig. 3 c; Jégu et al., 2004: 131, fig. 7 c). The difficulty in establishing homology among distict teeth rows in the premaxillary has been repeatedly discussed in the literature (Howes, 1982: 2; Vari & Harold, 2001: 12; Zanata & Vari, 2005: 32 – 33; Mirande, 2010: 420 – 421). Mirande (2010: 420) suggested that developmental data could help in establishing the homologies of distinct premaxillary teeth rows, since he supposed that, while most of the premaxillary teeth developed intraosseously (i. e., in bone crypts), some of them developed extraosseouly, although there is some difficulty in establishing whether the development of some teeth are intraosseous or extraosseous in the group (Trapani, 2001; Trapani et al., 2005). In our opinion, adequate hypothesis of homology between premaxilary teeth rows across the characifoms will ultimately depend on the identification and study of taxa presenting intermediate, transitional teeth arrangements between the main known types of teeth arrangements, whether using extant (Ribeiro et al., 2004) or extinct (Cione et al., 2009) examples. 2. Continuity of orbital ring. The series of ossifications surrounding the orbit is formed by six infraorbital bones, plus the supraorbital and the antorbital. The most common condition found among characiforms is to possess an incomplete orbital ring, either due to lack of contact between the sixth infraorbital with the supraorbital, and / or the supraorbital with the antorbital (e. g., Distichodontidae and Citharinidae; Vari, 1979: 296 – 299; Serrasalmidae; Machado-Allison, 1983: 163 – 164, 192, fig. 11; some Alestidae: Zanata & Vari, 2005: 6, fig. 1 B; Acestrorhynchus; Toledo-Piza, 2007: 713, fig. 14), or for the lack of supraorbital (e. g., Erythrinidae, Lebiasinidae, some Alestidae, and most Crenuchidae and Characidae; e. g., Weitzman & Fink, 1983; Vari, 1995; Buckup, 1993, 1998; Malabarba & Weitzman, 2003; Zanata & Vari, 2005). In those cases, the orbit is partially delimited by the frontal bone. A complete, continuous orbital ring is reported among characiforms in some Hemiodontidae (Roberts, 1974: 418 – 419), Hepsetidae (Roberts, 1969; Vari, 1995), all Ctenoluciidae (Roberts, 1969; Vari, 1995), some Alestidae (Zanata & Vari, 2005: 9), and, among Characidae, all Cynodontinae (e. g., Hydrolycus: Toledo-Piza, 2000: fig. 2), Agoniates (e. g, Géry, 1962: fig. 3), Salminus (e. g., Roberts, 1969; Machado-Allison, 1983: 192, fig. 11), Triportheus, and most Brycon species. Lignobrycon appears to be variable on this regard; a cleared and stained specimen examined from the rio de Contas basin (MZUSP 75230) possess a continuous orbital ring, while specimens from the rio de Una basin the sixth infraorbital does not contact the supraorbital (Castro & Vari, 1990: fig. 2; A. M. Zanata, pers. comm.). As earlier remarked by Zanata (2000: 28), species of Brycon typically possess a continuous orbital ring, but there are some exceptions. Zanata (2000: 28) reported that Brycon henni (a transandean Brycon species widely distributed at the Río Cauca and other river drainages of northeastern Colombia; Maldonado-Ocampo et al., 2005), and B. coxeyi, lack a contact between the sixth infraorbital and the supraorbital bones. During the course of the present study, we noticed that Brycon stolzmanni also lack a contact between these bones as well. Since all these species also share the presence of a dark patch of dark pigmentation at the opercle, the lack of continuity in the orbital ring shared by them is probably best interpreted as a derived condition within the genus.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F71FFF64EA4F9C4FB4AFA7A.taxon	materials_examined	3. Inner symphyseal dentary teeth pair. The presence of a inner symphyseal teeth pair, slightly smaller than the teeth from the main dentary series situated immediately ahead, is an uncommon feature among characiforms, being present in the serrasalmids Colossoma, Piaractus, and Mylossoma (Machado-Allison, 1986: 47), Mylesinus paucisquamatus (Jégu & Santos, 1988), Tometes (Jégu, Santos, Keith & Le Bail, 2002; Jégu, Santos & Belmont- Jégu, 2002), Utiaritichthys (Jégu et al., 1992), Myloplus (Jégu et al., 2003; Jégu et al., 2004), Myleus (Jégu & Santos, 2002), and in most alestids (Zanata & Vari, 2005: 46 – 47). Among characids, it is reported only for the representatives of Bryconinae, i. e., Brycon, Triportheus (Howes, 1982: 1 – 2), Henochilus (Castro et al., 2004: 501), and Chilobrycon (Géry & de Rham, 1981: 10). Contrary to Machado-Allison (1983: 174 – 175), Zanata & Toledo- Piza (2004: 105 – 106), Zanata & Vari (2005: 46 – 47), and Mirande (2010: 425), and as Uj (1990: 52 – 53), we prefer to consider the symphyseal teeth pair as a distinct, independent morphological character from the row of small inner row of conical teeth on the lingual crest of the teeth replacement trench. The symphyseal teeth pair has an intraosseous origin, being formed in an autonomous tooth fossa (cf. Roberts, 1967: 246, fig. 3; Machado-Allison, 1986: 47), unlike the inner conical teeth row, which has an extraosseous origin, being developed directly in the oral epithelium. Also, with the exception of the genera Brycon, Henochilus, Chilobrycon, and Chalceus, all remaining taxa displaying a symphyseal teeth lack an inner dentary teeth row. Mirande (2010: 425) states that Triportheus also possess a inner row of dentary teeth., apparently because he considered the small, distalmost teeth on the dentary as constituting the inner teeth row, since there is a overlap between the last teeth of the main dentary series and the first small teeth of dentary in the specimen of Triportheus nematurus he examined (Mirande, 2010: 419, fig. 60). This overlap is, however, apparently atypical, as it was not observed in the material of Triportheus herein examined (see Comparative material examined at Appendix 1). Although it makes sense for developmental reasons to consider the small dentary teeth as being homologous to the inner teeth row rather than to the main dentary teeth row since they are developed extraosseously as well (Trapani et al., 2005: 528; pers. obs.), the condition observed in Triportheus is not similar to the one found in Brycon, Chalceus, Brycon, and Henochilus, and it is questionable to consider it equivalent to the condition found in these latter genera. Contrary to the inner symphyseal dentary teeth, the presence of an inner dentary teeth row is a relatively widespread feature among characiforms. The presence of the inner symphyseal teeth is variable instraspecifically both in some alestids (Zanata & Vari, 2005: 47) and serrasalmids (e. g., Tometes makue; Jégu, Santos & Belmont-Jégu, 2002: 258, 260). In Brycon, the symphyseal teeth may be missing in large specimens of several of the large-sized species, such as Brycon vermelha, B. amazonicus, or B. falcatus, but is always present in juvenile and middle-sized specimens of all species.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F71FFF64EA4F9C4FB4AFA7A.taxon	discussion	Remarks. The genus Brycon was originally proposed by Müller & Troschel (1844) to include some species previously assigned to the genus Chalceus Cuvier (C. macrolepidotus, C. amazonicus, and C. opalinus), plus three species that were described in that contribution, Brycon falcatus, B. schomburgkii, and B. pesu. Müller & Troschel (1844), however, included in Brycon the type species of Chalceus, C. macrolepidotus, a mistake that prompted Valenciennes (in Cuvier & Valenciennes, 1850: 239) and Kner (1859: 9) into not recognizing Brycon as a valid genus. Günther (1864: 333 – 334) provided a new diagnosis for Brycon, and ever since this genus has been recognized as valid. Brycon, as currently understood, comprises seven synonyms, most of which were never consistently used in the literature. Two of these synonyms, Chalcinopsis Kner (1858: 223) and Othonophanes Eigenmann (1903: 145) were described having as type-species taxa from Central America or trans-andean South America, and are not further discussed here. Braum (1983 b) discussed the synonym of Othonophanes with Brycon. A third synonym, Holobrycon Eigenmann (1909: 33), was proposed to include Brycon pesu, a species not treated in the present contribution. See Howes (1982) for the synonymization of Holobrycon into Brycon.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F71FFF64EA4F9C4FB4AFA7A.taxon	description	So far, four phylogenetic analyses has been undertaken aiming to understand the intrarelationships of the genus. Hilsdorf et al. (2008) presented a hypothesis of relationships within the genus using partial 16 S rDNA mitochondrial sequences of 13 Brycon species, plus the Bryconinae Henochilus wheatlandii. A more comprehensive analysis of the phylogenetic relationships of the genus, based on morphological data was undertaken by Zanata (2000), but still remains unpublished. Both hypotheses agree in identifying a well-defined, derived group that includes the large-sized, diverse-colored Brycon species widespread in the large floodplain rivers of cis-andean South America that includes B. falcatus, B. orbignyanus, B. hilarii, B. orthotaenia, B. amazonicus, B. whitei, and B. melanopterus, and which roughly corresponds to the Brycon falcatus - group of Howes (1982). Hilsdorf et al. (2008) hypothesis identifies monophyletic clades including the Brycon species of eastern Brazil (B. opalinus, B. insignis, B. ferox, and B. nattereri) plus Henochilus wheatlandii, and a monophyletic clade including some trans-andean species. This roughly agrees with the hypothesis of Zanata (2000), though in her hypothesis the Brycon species from eastern Brazil does not form a monophyletic clade. Recently, two additional molecular phylogenies were presented for the genus. The first, by Abe et al. (2014) presented a more comprehensive hypothesis of Brycon (including Salminus), based on partial sequences of two mitochondrial and three nuclear genes. Abe et al. (2014) analysed 19 out of the currently 44 valid species of Brycon, plus the monotypic genera Chilobrycon and Henochilus, and all species of Salminus, and considered Brycon in the present sense as paraphyletic without the inclusion of Salminus. They recovered five monophyletic clades within Brycon, the most basal one including four of the analysed trans-andean Brycon species, plus Chilobrycon deuterodon, and then successively a clade including all Salminus species, a clade formed solely by Brycon pesu, and a clade formed by Brycon amazonicus, B. hilarii, B. gouldingi, B. orbignyanus, B. orthotaenia, and B. moorei, which is sister of a clade containing two subclades, one composed by B. falcatus and B. melanopterus, and the other including exclusively Brycon species from eastern Brazil, as well as Henochilus wheatlandii. Most of these clades seems to concur with morphological evidence, but the inclusion of Salminus within Brycon, as well as the relationships among them, are apparently at odds with the morphological evidence. The second, by Travenzoli et al. (2015), focused on the species occurring in eastern Brazil and provided additional evidence for the monophyly of a clade composed by Brycon species occurring in eastern Brazil, including Henochilus wheatlandii. The position of Henochilus wheatlandii not only within Brycon, but also within the clade of Brycon from eastern Brazil, already found by Hilsdorf et al. (2008) and confirmed by Abe et al. (2014) and Travenzoli et al. (2015), is revealing. The other monotypic genus of Bryconinae, Chilobrycon, was also recovered within the genus Brycon, both using morphological (Zanata, 2000) and molecular (Abe et al., 2014) data. It seems likely that both genera are synonyms of Brycon, but until both a comprehensive morphological study and a broader molecular analysis of the genus are undertaken, no changes at the generic level within Bryconinae seem advisable.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F71FFF64EA4F9C4FB4AFA7A.taxon	distribution	Distribution. From ríos Grijalva and Usumancita, Chiapas state, Mexico (Miller et al., 2005) southwards to Panama in Central America (Hildebrand, 1938), and transandean river systems, from Río Jequetepeque in northern Peru (Pearson, 1937) and Río Tumbes at the peruvian / ecuatorian border (Chirichigno, 1963; Géry & de Rham, 1981) (but actually occurring southward down to Río Santa, Departamento Ancash, central-northern coastal Peru; H. Ortega, pers. comm ..; Brycon cf. atrocaudatus; MUSM 2990, MUSM 3003) to the Lake of Maracaibo system, Venezuela (Moscó Morales, 1988). At cis-andean South America, the genus Brycon is widespread across the major river systems, i. e., Río Orinoco, rio Amazonas, Rio de la Plata, and rio São Francisco basins, guyanese river systems (from Essequibo River in Guyana to Mana River in French Guiana), and small coastal river systems from the rio São Francisco mouth southwards to the rio Paraíba do Sul basin in eastern Brazil. There are no records, and the genus is with all likelihood absent, from the several rivers drainages from northeastern Brazil between the mouths of the rio São Francisco and rio Capim, and from the coastal river systems from southeastern South America situated to the south from the Baía de Guanabara in Rio de Janeiro, eastern Brazil.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F79FFE84EA4FA37FBA0FD10.taxon	discussion	The key provided below summarizes the main diagnostic features to diagnose the different cis-andean Brycon species. As is always the case with a dichotomous key, care should be exercised in its use, and the “ Diagnosis ” section of each species should be checked for additional diagnostic features and detailed comparisons. Juveniles (<100 mm SL) of snout-pointed Brycon species from eastern Brazil (steps 4 to 8) cannot be reliably distinguished among themselves. Brycon amazonicus populations from the Amazon basin at Peru possess a similar color pattern to B. hilarii, and the diagnosis of both species in the latter area is difficult (see under Brycon amazonicus).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F67FFEC4EA4FAF9FCB0F9EF.taxon	diagnosis	Diagnosis. Brycon stolzmanni can be distinguished from all remaining cis-andean Brycon species, except B. coxeyi, by possessing a distinct patch of dark pigmentation on the opercle (vs. patch of dark pigmentation on the opercle absent in the remaining cis-andean Brycon species) and infraorbital series not closed, with the orbital margin partially delimited by the frontal bone (vs. infraorbital series closed, frontal excluded from the orbital series by the contact between the supraorbital and the sixth infraorbital bone). Brycon stolzmanni can be distinguished from B. coxeyi by possessing the dark patch of pigmentation on the opercle diffuse and covering the upper half of the bone (vs. dark patch of pigmentation on opercle very conspicuous and typically lying at its center).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F67FFEC4EA4FAF9FCB0F9EF.taxon	description	Description. Morphometric data are presented in Table 1. Relatively small-sized species, largest examined specimen 214.0 mm SL. Body moderately high. Largest body height at dorsal-fin level. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly concave to slightly convex from latter point to basis of supraoccipital process, and moderately convex from latter point to dorsal-fin origin, straight along dorsal-fin basis, straight to slightly convex from dorsal-fin terminus to adipose-fin origin. Dorsal profile of caudal peduncle slightly concave. Ventral profile slightly convex from lower lip to pelvic-fin insertion, slightly convex to straight from this point to anal-fin origin and approximately straight along anal-fin base. Ventral profile of caudal peduncle slightly concave. Head profile rounded to slightly acute anteriorly, mouth terminal. Maxillary extending posteriorly to vertical slightly anterior to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Five (7), 6 (16), 7 (10) or 8 (1) tricuspidate teeth in outer series. Two (4), 3 (25), or 4 (7) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 3 (30) or 4 (6) tricuspidate teeth between the first and third rows. Two teeth in third premaxillary row, medial teeth largest, symphyseal teeth smaller, both pentacuspidate. Maxillary with distal portion expanded and rounded in profile. Eleven to 18 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 6 (1), 7 (3), 8 (1), 9 (1), or 10 (1) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, tetra- to pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, tri- to unicuspidate. Inner (lingual) series consisting of a single unicuspid symphyseal tooth, considerably smaller than teeth of main series anterior to it, plus row of 5 small, unicuspidate teeth, originating at level of 7 th tooth of main series. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-three (1), 44 (3), 45 (6), 46 (10), 47 (12), 48 (3), or 49 (1) scales in lateral line series; laterosensory tube simple, tube often deflected downwards. Horizontal scale rows between dorsal-fin origin and lateral line 8 (17), 9 (16), or 10 (3). Horizontal scale rows between lateral line and pelvic-fin 4 (4), 5 (15), or 6 (17). Circumpeduncular scales 16 (18), 17 (9), or 18 (9). Dorsal-fin rays ii, 9. Dorsal fin origin slightly anterior to middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 12 th vertebrae. Dorsal fin displaying numerous (c. 10 – 35 per fin-ray main branch) small hooks on last unbranched and posterior main branch of all branched rays in one specimen (ROM 55348, 128.8 mm SL). Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 19 (1), 20 (6), 21 (16), 22 (12), or 23 (1). First anal-fin pterygiophore inserting behind haemal spine of 22 th vertebrae. Last unbranched and anterior 2 – 3 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 15 – 35 per fin-ray main branch) small hooks associated with dense, gelatinous tissue on last unbranched and posterior main branch of all branched rays except posteriomost ray in one specimen (ROM 55348, 128.8 mm SL). Small hooks also present in the anterior main anal-fin branch in the two anteriormost anal-fin rays. Sheath of scales composed of a single series composed by 16 – 20 rectangular scales, covering basis of anal-fin rays. Pectoral-fin rays i, 11 (1), 12 (15), 13 (18), or 14 (2). Pelvic-fin rays i, 7. Branched pelvic-fin rays with minutes hooks (c. 10 per branch) on distal portion of posterior ray branches in one specimen (ROM 55348, 128.8 mm SL). Main caudal-fin rays 10 / 9. Caudal fin forked, lobes rounded. Four branchiostegal rays. First gill arch with 11 (1) lower gill rakers, 11 (1) upper gill rakers and 1 at angle. Vertebrae 43 (1). Supraneurals 9 (1). Coloration in alcohol. Top of head, snout, supraorbital, sixth and dorsal portion of fifth infraorbitals, and dorsal portion of body dark grey to dark brown. Dentary, maxillary, lower infraorbitals, gular area and ventral portion of body light to cream-colored. Opercle dark-colored, its lower half dark-grey, suffused with silvery pigmentation, while upper half dark brown. Humeral blotch present, split in two halves, lower half more conspicuous, roughly triangular-shaped, with the apex directed downwards, crossed at its lower portion by the lateral line, extending horizontally from the third to the fifth to sixth lateral line scales. Upper half of humeral blotch less conspicuous, separated from lower half of humeral blotch by a narrow clear-colored area, two and a half scales rows high. Caudal peduncle blotch present, relatively conspicuous, rounded, extending throughout 3 – 4 last lateral line scales into basis of 3 – 4 innermost caudal-fin rays. Caudal, anal and dorsal-fins with numerous small dark chromatophores scattered over interradial membranes. Pectoral and pelvic-fins with relatively few dark chromatophores on interradial membranes. Adipose fin dark brown, with a conspicuous clear-colored rounded spot at its basis. Sexual dimorphism. The single examined specimen bearing hooks on dorsal, pelvic- and anal-fins (ROM 55348, 128.8 mm SL) was found to be a male upon dissection. The largest specimen examined, NMW 62728 (214.0 mm SL), is a female with well-developed ovaries, which does not possess fin hooks.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F67FFEC4EA4FAF9FCB0F9EF.taxon	distribution	Distribution. Brycon stolzmanni is known from the upper Río Marañon and its tributaries at the Departamentos Cajamarca, Lambayeque, and Amazonas, Peru (Fig. 10). Fowler (1943: 4) and Howes (1982: 44) incorrectly assigned the species for the Pacific slope of the Andean cordillera. Ecological notes. Brycon stolzmanni is recorded from Andean rivers in altitudes ranging at least from 870 – 1087 meters high. The type locality, Chota, is situated in the headwaters of the Río Chamaya, a tributary of the upper Río Marañon, at a considerably higher altitude (2400 meters high). However, its is uncertain whether the type specimens were collected at Chota itself or in a neighboring river valley, and consequently we did not included this record in the altitudinal range cited above. The Río Marañon and its tributaries at the area of occurrence of Brycon stolzmanni are swift-flowing, white-water rivers possessing coarse gravel and rocky substrates running over braided channels, deeply entrenched in valleys crossing the mountains.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F67FFEC4EA4FAF9FCB0F9EF.taxon	discussion	Remarks. Steindachner (1879 a: 152) succinctly described Brycon stolzmanni, a complete description and a illustration of the species appearing a little later (1879 b: 22). The species was described based on two specimens collected at Chota, upper Río Marañon basin, Departamento Cajamarca, Peru. The two syntypes (NMW 62731) were examined, the largest of which is herein designated as the lectotype of the species. There have been almost no subsequent published information on the species, and in fact, Pearson (1937) report of the species for several localities in the upper Río Marañon basin in Peru was the sole paper to add original information on the species up to now.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F67FFEC4EA4FAF9FCB0F9EF.taxon	materials_examined	Material examined. Type material. NMW 62731: 1 (1, 124.5 mm SL): Chota (Peru, Depto. Cajamarca, Rio Marañon basin, c. 6 ° 33 ’ S, 78 ° 39 ’ W); Stolzmann, no date. Lectotype by present designation of Brycon stolzmanni Steindachner. NMW 62731: 2 (1, 88.4 mm SL); same data as lectotype. Paralectotype. Non types. All from Peru, Rio Maranõn drainage. Departamento Cajamarca: ROM 52237 (5 of 51, 146.8 – 160.5 mm SL), approx. 74 km W of road going N to Jaen, between Pucara and Guabal, Chamaya River, Huacabamba River (tributary Amazon / Marañon drainage), c. 5 ° 57 ’ S, 79 ° 13 ’ W; E. Holm & J. Patalas, 8 Jul 1986. CAS 77371 (69, 27.3 – 72.5 mm SL), Rio Crisnejas, at mouth of Rio Paipay at Paipay, c. 7 ° 21 ’ S, 77 ° 53 ’ W; N. E. Pearson, Aug 1923. NMW 62728 (1, 214.0 mm SL); NMW 62933 (1, 88.5 mm SL), Celendín, Hacienda Callacate, 6 ° 46 ' S, 78 ° 16 ' W; W. Taczanowski, 1880. MUSM 41333 (9, 20.5 – 37.2 mm SL): Celendín, Río Llanguat (Río Marañon basin), 6 ° 47 ’ 19 ’’ S, 78 ° 12 ’ 55 ’’ W; H. Ortega et al., 25 Sept 2011. Departamento Amazonas: ROM 55347 (1, 163.7 mm SL), Luya, Huanabamba, about 16 km by road upstream from Balsas, Amazon / Marañon drainage, 6 ° 52 ' S, 78 ° 00 ' W; E. Holm et al., 27 June 1986. ROM 55348 (2, 128.8 – 132.7 mm SL), confluence of “ El Río ” (creek running through Balsas) at Puerto Balsas, 6 ° 50 ' S, 78 ° 01 ' W; E. Holm et al., 28 Jun 1986. UMMZ 185306 (28, 49.6 – 130.7 mm SL); UMMZ 216716 (5, 107.3 – 185.5 mm SL): Rio Maranõn at Balsas, 6 ° 50 ’ S, 78 ° 1 ’ W; N. E. Pearson, July 1923. CAS 77370 (ex IU 17599) (6 of 94, 20.9 – 33.2 mm SL), Río Marañon, Pusoc (Guayabamba), above Balsas, c. 6 ° 57 ’ S, 78 ° 00 ’ W; N. E. Pearson, Aug 1923. CAS 77372 (32, 21.5 – 42.9 mm SL), Río Marañon at Tingo de Pauca, mouth of Río Crisnejas, c. 7 ° 20 ’ S, 77 ° 49 ’ W; N. E. Pearson, Sept 1923. MUSM 10611 (5, 65.8 – 94.0 mm SL): Río Uctubamba, Lluhuana; F. Chang, 23 Nov 1996. MUSM 40548 (1, 85.0 mm SL): Bongará, Shipasbamba, Río Uctubamba (trib. Río Marañon), 5 ° 54 ’ 45 ’’ S, 78 ° 3 ’ 50 ’’ W; M. Jaico et al., 12 Jul 2011. MUSM 41145 (44, 45.9 – 105.2 mm SL); ZUEC 6850 (4, 58.9 – 101.4 mm SL): Amazonas, Bongara, Jazan, Río Tingo (trib. Río Uctubamba), 5 ° 55 ’ 2 ’’ S, 78 ° 4 ’ 35 ’’ W; M. Hidalgo & I. Corahua, 29 June 2011. MUSM 41146 (8, 62.1 – 99.0 mm SL): Amazonas, Bongara, Jazan, Río Utcubamba (island), 5 ° 55 ’ 1 ’’ S, 78 ° 4 ’ 32 ’’ W; M. Hidalgo & I. Corahua, 29 June 2011. Departamento Lambayeque: MUSM 30230 (1, 35.5 mm SL): Ferreñafe, Kañaris, CCA Huancabamba, Río Cañariaco, mouth at Río Huancabamba, 5 ° 56 ’ 15 ’’ S, 79 ° 15 ’ 47 ’’ W; B. Renjifo & V. Meza, 24 May 2007. Without data: MUSM uncat. (1, 133.7 mm SL, cs).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F63FFE24EA4F99BFE0EFA03.taxon	description	Topo, a tributary of Río Paztaza, 1 ° 25 ’ S, 78 ° 11 ’ W ”).? Brycon species: Géry, 1972 b: 7 – 8 (Río Villano, tributary of Río Cururay, Amazon basin).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F63FFE24EA4F99BFE0EFA03.taxon	diagnosis	Diagnosis. Brycon coxeyi can be distinguished from all remaining cis-andean Brycon species, except Brycon stolzmanni, by possessing a distinct patch of dark pigmentation on the opercle (vs. patch of dark pigmentation on the opercle absent in the remaining cis-andean Brycon species) and infraorbital series not closed, with the orbital margin partially delimited by the frontal bone (vs. infraorbital series closed, frontal excluded from the orbital series by the contact between the supraorbital and the sixth infraorbital bone). Brycon coxeyi can be distinguished from Brycon stolzmanni by possessing the dark patch of pigmentation on opercle well-defined, typically lying at its center (vs. dark patch of pigmentation diffuse and restricted to the upper half of opercle in B. stolzmanni).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F63FFE24EA4F99BFE0EFA03.taxon	description	Description. Morphometric data are presented in Table 2. Relatively small-sized species, largest examined specimen 225.0 mm SL. Body 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, slightly concave from latter point to basis of supraoccipital process, and moderately convex from latter point to dorsal-fin origin, straight along dorsal-fin basis, straight to slightly convex from dorsal-fin terminus 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. Maxillary extending posteriorly to vertical slightly anterior to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Five (2), 6 (24), 7 (25), 8 (5), or 9 (1) tricuspidate teeth in outer series. Two (4), 3 (42), or 4 (7) tri- to tetracuspidate teeth in second, inner premaxillary row, plus 2 (4), 3 (42), or 4 (7) tricuspidate teeth between the first and third rows. Two teeth in third premaxillary row, medial teeth largest, symphyseal teeth smaller, both pentacuspidate. Maxillary with distal portion expanded and rounded in profile. Nine to 21 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 6 (2), 7 (1), 9 (1), 11 (1), or 13 (1) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, tetra- to pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, tri- to unicuspidate. Inner (lingual) series consisting of a single unicuspid symphyseal tooth, considerably smaller than teeth of main series anterior to it, plus row of 6 small, unicuspidate teeth, originating behind last tooth of the main series. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-two (1), 43 (1), 44 (3), 45 (15), 46 (17), 47 (8), 48 (9), 49 (2), 50 (1), or 52 (1) scales in lateral line series; laterosensory tube simple, straight or slightly deflected downwards. Horizontal scale rows between dorsal-fin origin and lateral line 8 (22), 9 (34), or 10 (2). Horizontal scale rows between lateral line and pelvic-fin 4 (9), 5 (43), or 6 (5). Circumpeduncular scales 14 (4), 15 (4), 16 (30), 17 (16), 18 (1), or 19 (1). Dorsal-fin rays ii, 9, one specimen ii, 10. Dorsal fin origin at middle of SL. Dorsal fin displaying numerous (c. 15 – 30 per fin-ray main branch) small hooks on last unbranched and all branches (more numerous on last branch) of all branched rays in 13 specimens (MEPN 11101, 205.0 mm SL; MEPN 11108, 155.4 mm SL; MEPN 11115, 144.8 mm SL; MEPN 11109, 150.9 mm SL; MEPN 11103, 187.0 mm SL; MEPN 11105, 168.1 mm SL; MEPN 11114, 2, 153.0 – 157.2 mm SL; MEPN 11112, 122.6 mm SL; MUSM 6127, 2, 126.6 – 135.4 mm SL; MUSM 6136, 2, 131.5 – 135.8 mm SL). Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 21 (6), 22 (17), 23 (33), or 24 (10). Last unbranched and anterior 2 – 3 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 20 – 30 per fin-ray main branch) small hooks associated with dense, gelatinous tissue on last unbranched and posterior main branch of all branched rays except posteriomost ray in 15 specimens (MEPN 11101, 205.0 mm SL; MEPN 11108, 155.4 mm SL; MEPN 11115, 144.8 mm SL; MEPN 11109, 2, 149.6 – 150.9 mm SL; MEPN 11103, 187.0 mm SL; MEPN 11105, 168.1 mm SL; MEPN 11102, 218.0 mm SL; MEPN 11114, 2, 153.0 – 157.2 mm SL; MEPN 11112, 122.6 mm SL; MUSM 6127, 2, 126.6 – 135.4 mm SL; MUSM 6136, 2, 131.5 – 135.8 mm SL). Small hooks also present in the anterior main anal-fin branch in the two anteriormost anal-fin rays. Sheath of scales composed of a single series composed by 17 – 20 rectangular scales, covering basis of anal-fin rays. Pectoral-fin rays i, 12 (28), 13 (24), or 14 (4). Pelvic-fin rays i, 7, one specimen i, 6, and another single specimen i, 8. Branched pelvic-fin rays with minutes hooks (c. 10 per branch) on distal portion of posterior ray branches in 14 specimens (MEPN 11101, 205.0 mm SL; MEPN 11108, 155.4 mm SL; MEPN 11115, 144.8 mm SL; MEPN 11109, 2, 149.6 – 150.9 mm SL; MEPN 11103, 187.0 mm SL; MEPN 11105, 168.1 mm SL; MEPN 11102, 218.0 mm SL; MEPN 11114, 2, 153.0 – 157.2 mm SL; MEPN 11112, 122.6 mm SL; MUSM 6127, 2, 126.6 – 135.4 mm SL; MUSM 6136, 2, 131.5 – 135.8 mm SL). Main caudal-fin rays 10 / 9. Caudal fin forked, lobes rounded. Four branchiostegal rays. First gill arch with 10 (1), 11 (2), or 14 (1) lower gill rakers, 9 (3) or 10 (1) upper gill rakers and 1 at angle. Vertebrae 44 (1). Coloration in alcohol. Top of head, snout, supraorbital, first, sixth, and fifth infraorbital bones, and dorsal portion of body dark grey to dark brown. Dentary, maxillary, gular area, and ventral portion of body cream-colored. Second through fourth infraorbital bones silvery in specimens retaining guanine, with a brownish tinge. Opercle silvery, with a large, rounded conspicuous dark patch of pigmentation typically lying at its central area, with the exception of darkened large specimens, described below. Humeral blotch present, little conspicuous, more evident in smaller specimens or specimens lacking scales on humeral region, split in two halves. Humeral blotch vertically elongated, lower half more conspicuous, crossed at its middle by the lateral line, extending horizontally from the third to the fourth lateral line scales. Upper half of humeral blotch less conspicuous, separated from lower half of humeral blotch by a narrow clear-colored area, two scales rows high. Lateral portions of body clear, with a silvery hue in specimens retaining guanine. Body with multiple moderately conspicuous longitudinal rows of wavy dark stripes, formed by pigmentation concentrated along scale junctions, more discernible dorsally. Caudal peduncle blotch present, moderately to highly conspicuous, rounded, extending throughout 4 – 5 last lateral line scales into basis of 5 – 6 innermost caudal-fin rays. Caudal, anal and dorsal-fins with numerous small dark chromatophores scattered over interradial membranes. Pectoral and pelvic-fins with relatively few dark chromatophores on interradial membranes. Adipose fin light grey, with a well-defined clear area at its basis. Diffuse dark pigmentantion present at lateral portion of body in several middle to large sized specimens (120 – 220 mm SL, e. g., MEPN 11101, 1, 205.0 mm SL, MEPN 11112, 122.6 mm SL). Some specimens (e. g., MEPN 11115, 3, 164.6 – 168.0 mm SL; MEPN 11105, 2, 168.1 – 173.4 mm SL; MEPN 11103, 187.0 mm SL; MUSM 6141, 145.1 mm SL; MUSM 6136, 3, 131.5 – 135.8 mm SL) develop a intense dark pigmentantion on body sides, presenting a much enlarged dark blotch on opercle, covering the entire middle and lower surface of that bone, and sides of body with a marbled pattern, with intense dark and white / silvery pigmentation alternating (see item “ Sexual Dimorphism ”, below). A single specimen (MEPN 11103, 187.0 mm SL) with a large amount of dark pigmentation on the third to the fifth infraorbital bones. Color in life. Description based on pictures of several freshly-collected specimens from the Río Machinaza (upper Río Zamora / Río Santiago basin), taken by R. Barriga and collaborators. Snout, top of head, and dorsum plumbeous-grey to brown; sides of body silvery, with a plumbeous tinge. Lower portion of head and branchiostegal rays clear, tinged with orangish in larger specimens. Dark patch of pigmentation in opercle and caudal-peduncle blotch conspicuous. Longitudinal wavy stripes moderately conspicuous, more discernible dorsally. Pectoral and pelvic-fins reddish. Anal and caudal fin with red pigmentation concentrated at its basis. Sexual dimorphism. Hooks were observed at the anal-fin rays (and in most cases, also dorsal-fin rays and distal portion of pelvic-fin rays) of 18 specimens (MEPN 11101, 205.0 mm SL; MEPN 11108, 155.4 mm SL; MEPN 11115, 144.8 mm SL; MEPN 11109, 2, 149.6 – 150.9 mm SL; MEPN 11103, 187.0 mm SL; MEPN 11105, 168.1 mm SL; MEPN 11102, 218.0 mm SL; MEPN 11114, 2, 153.0 – 157.2 mm SL; MEPN 11112, 122.6 mm SL; MUSM 6141, 145.1 mm SL; MUSM 6127, 2, 126.6 – 135.4 mm SL; MUSM 6144, 108.2 mm SL; MUSM 6136, 3, 131.5 – 135.8 mm SL). Specimen MEPN 11115 (144.8 mm SL) was dissected and is a male, with well-developed testes. On other hand, a specimen lacking any hooks but collected alongside two specimens possessing hooks, MEPN 11114 (182.0 mm SL), proved to be female on dissection, with ripe oocytes. A moderate to intense dark pigmentation on body sides and a enlarged dark blotch on opercle are present in some specimens and seem to be an additional dimorphic feature, since most specimens presenting these color features also present hooks on fins. However, some specimens presenting an enlarged dark blotch on opercle (MEPN 11115, 3, 164.6 – 168.0 mm SL; MEPN 11117, 2, 147.9 – 177.1 mm SL; MUSM 6144, 1, 108.2 mm SL) does not possess any fin hooks, and conversely, some specimens presenting hooks (MEPN 11115, 144.8 mm SL; MUSM 6127, 2, 126.6 – 135.4 mm SL) present instead the “ normal ” color pattern. Brycon guatemalensis is known to possess mature males with a marbled darkened pattern on sides (e. g., Bussing, 1987) and males of Brycon species from all river systems of Pacific versant of the Andes in Ecuador also display a darkened color pattern when mature (R. Barriga, pers. comm ..). Common names. “ Blanco ”, “ sábalo ” (Zamora-Chinchipe, Ecuador; R. Barriga, pers. comm.).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F63FFE24EA4F99BFE0EFA03.taxon	distribution	Distribution. Brycon coxeyi is only known from upper tributaries of the Río Marañon basin in Ecuador and Peru, at altitudes ranging from 500 – 1392 meters a. s. l. (Fig. 10). Ecological notes. Brycon coxeyi is found in fast-running, rocky-bottomed rivers of the Andean piedmont in Ecuador. Two of the the rivers known to harbor Brycon coxeyi populations, the Río Nangaritza and the Río Machinaza, both tributaries of the upper Río Zamora / Río Santiago basin, are fast-running, rocky-bottomed blackwater rivers (R. Barriga, pers. comm ..). The Río Comainas in Cordillera del Condor, Peru, where the species also occurs, is a shallow (0.3 – 1.9 meters deep), relatively small (8 – 20 meters wide) clear-water river, with a rocky / gravel / sandy / clay substrate from most of the year, except after heavy rains, when it becomes torrential, with turbid waters (Vari et al., 1995: 294). Crushed vegetal matter (leaves and apparently also fruit remains) were found in two specimens (MEPN 11115, 163.1 mm SL; MEPN 11114, 182.0 mm SL) examined for gut contents. Specimens presenting fin hooks and as such presumably in breeding period were found in March, April, July, September, and November. A female with ripe oocytes (MEPN 11114, 182.0 mm SL) was collected in April and a male with ripe testes (MEPN 11115, 144.8 mm SL) was collected in September. Specimens with a large amount of fat in the abdominal cavitiy were collected in February (MEPN 11119).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F63FFE24EA4F99BFE0EFA03.taxon	discussion	Remarks. Fowler (1943: 3) described Brycon coxeyi based on a single specimen collected at the Río Topo in the upper Río Paztaza basin, Ecuador. He noticed that the species was “ apparently not closely related to any of the many known forms distributed in the Atlantic drainage of the eastern Andes ”, and compared it with to Brycon henni Eigenmann, a species occurring in several trans-andean river drainages of Colombia, and B. stolzmanni. Fowler (1943) incorrectly assumed that the latter could be differentiated from Brycon coxeyi by supposedly lacking the opercular dark patch of pigmentation, when in fact B. stolzmanni possess it. Both species can be differentiated by the characters mentioned in the item “ Diagnosis ”, above. The holotype (ANSP 70156) is in a very poor condition of preservation, being very soft, with no scales left, most fins broken and the opercle bone almost disarticulated from the rest of the opercular apparatus (see Fig. 11). Still, it displays the rounded dark patch of pigmentation which is characteristic from the remaining Brycon specimens from the upland areas of the Amazon basin in Ecuador herein assigned to the species. As noticed by Böhlke (1958: 24 – 25) and Lima (2003: 175 – 176), the type-locality given by Fowler (1943) for Brycon coxeyi is incorrect, and was amended by the latter as “ Ecuador, Napo-Paztaza, Hacienda Mascota, Río Topo, a tributary of Río Paztaza, 1 ° 25 ’ S, 78 ° 11 ’ W ”. However, according to the old label present in the jar of the holotype of Brycon coxeyi, the specimen was not collected at the Río Topo itself but rather at the Río Paztaza near the mouth of the Río Topo, and should be again corrected, this time to “ Ecuador, Província Tungurahua, Hacienda Mascota, Río Paztaza, near mouth of Río Topo, c. 1 ° 25 ’ S, 78 ° 11 ’ W ”. Boulenger (1898 a: 3) examined seven Brycon specimens from the Río Mira and Río Peripa (Río Guayas basin), plus one specimen from the Río Zamora, all of which he identified as Brycon atricaudatus (= B. atrocaudatus). We have not re-examined the specimen from the Río Zamora (which is presumably deposited at the MSNG), but judging both from the locality and its meristic data (lateral line scales 46, scales above lateral line 9, and anal-fin rays 27 — i. e., iii, 24 — Boulenger, 1898 a: 3), this record is with all likelihood referable to Brycon coxeyi. Tortonese (1939: 49) re-examined this same specimen and incorrectly re-identified it as Brycon oligolepis. Géry (1972 b: 7 – 8) cited an unidentified species of Brycon based in specimens collected in the Río Curaray basin (not “ Cururay ”), a tributary of the Río Napo, Ecuador. Judging by the color pattern and locality data from these specimens, this record might refer to Brycon coxeyi, though anal-fin rays counts are too high (iv, 26 – 27, according to Géry, 1972: 7). Howes (1982: 21), without presenting any evidence to support such assertion, suggested that Brycon coxeyi might represent a synonym of B. melanopterus. However, both species are very distinct from each other and does not even seem to be closely related (see the account for Brycon melanopterus, below).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F63FFE24EA4F99BFE0EFA03.taxon	materials_examined	Material examined. Type material: ANSP 70156 (1, 131.6 mm SL): Ecuador, Província Tungurahua, Hacienda Mascota, Río Paztaza, near mouth of Río Topo, c. 1 ° 25 ’ S, 78 ° 11 ’ W; W. Judson Coxey, April 1931. Holotype of Brycon coxeyi Fowler. Non types. Ecuador: Província Zamora-Chinchipe, Río Zamora / Río Santiago basin: MEPN uncataloged (1, 148.5 mm SL): Río Nupatakaine, 500 m from Destacamento Shaine, at confluence with Río Nangaritza, c. 4 ° 28 ' S, 78 ° 39 ' W; R. Barriga, 27 July 1993. MEPN 17484 (1, 94.6 mm SL): Quebrada Mayayacu. 2 km from its mouth at Río Nangaritza, 3 ° 59 ’ 9 ’’ S, 78 ° 38 ’ 27 ’’ W; R. Barriga, 18 July 1993. MEPN uncataloged (1, 192.9 mm SL): Río Nangaritza, near sitio El Dorado, 3 ° 46 ' S, 78 ° 38 ' W; R. Barriga & M. Ibarra, 23 Sept 1978. MEPN 17494 (8, 50.4 – 123.6 mm SL); ZUEC 11325 (1, 106.3 mm SL): Quebrada Yunganza, trib. Río Santiago, 3 ° 24 ’ 00 ’’ S, 78 ° 33 ’ 25 ’’ W; R. Barriga, M. Ibarra & R. Loma, 27 March 1979. MEPN 11116 (1, 111.2 mm SL): Río Nangaritza; R. Barriga & M. Ibarra, 18 March 1979. MEPN 11114 (5, 153.8 – 182.0 mm SL): Río Machinaza, 3 ° 45 ’ 43 ’’ S, 78 ° 29 ’ 53 ’’ W; R. Barriga et al., 14 Apr 2009. MEPN 11115 (6, 152.3 – 167.6 mm SL): Río Machinaza, 3 ° 45 ’ 43 ’’ S, 78 ° 29 ’ 46 ’’ W; R. Barriga et al., 13 Sept 2009. MEPN 11102 (1, 218.0 mm SL): Zamora-Chinchipe, Río Machinaza Grande 0.5 km N of Garita de Aurelean, 3 ° 45 ’ 34 ’’ S, 78 ° 29 ’ 49 ’’ W; R. Barriga, J. Sandoval & J. Sharupi, 19 Nov 2008. MEPN 11103 (1, 187.0 mm SL): Zamora-Chinchipe, Río Machinaza, Peñas 3 ° 47 ' 1 '' S, 78 ° 29 ' 35 '' W; M. Andrade, 23 Jul 2010. MEPN 11104 (2, 149.5 – 179.8 mm SL): Zamora-Chinchipe, Río Machinaza, 500 m below confluence with Río La Zanza, 3 ° 45 ’ 46 ’’ S, 78 ° 29 ’ 46 ’’ W; R. Barriga & M. Andrade, 4 Oct 2009. MEPN 11105 (3, 168.1 – 180.9 mm SL): Zamora-Chinchipe, meeting of Río Zarza with Río Blanco, 3 ° 45 ’ 44 ’’ S, 78 ° 30 ’ 39 ’’ W; R. Barriga, J. Sandoval & S. Charupe, 9 Nov 2008. MEPN 11106 (4, 130.7 – 177.1 mm SL): Zamora-Chinchipe, Río Machinaza, 3 ° 45 ’ 21 ’’ S, 78 ° 29 ’ 34 ’’ W; R. Barriga et al., 13 Sept 2009. MEPN 11106 (4, 130.7 – 177.1 mm SL): Zamora-Chinchipe, Río Machinaza, 3 ° 45 ’ 21 ’’ S, 78 ° 29 ’ 34 ’’ W; R. Barriga et al., 13 Sept 2009. MEPN 11108 (4, 155.4 – 177.1 mm SL): Zamora-Chinchipe, Río Machinaza, 0,5 km above confluence with Río La Zalza, 3 ° 45 ’ 45 ’’ S, 78 ° 29 ’ 46 ’’ W; R. Barriga & M. Andrade, 4 Nov 2009. MEPN 11109 (13, 59.6 – 150.0 mm SL): Zamora- Chinchipe, Sachavaca Alta, Río Machinaza, 3 ° 41 ’ 13 ’’ S, 78 ° 29 ’ 27 ’’ W; M. Andrade, 29 – 30 Jul 2011. MEPN 11110 (4, 79.3 – 135.8 mm SL): Zamora-Chinchipe, 500 m above confluence of Río Zorza with Río Blanco, 3 ° 45 ’ 43 ’’ S, 78 ° 30 ’ 38 ’’ W; R. Barriga, J. Sharupe & J. Sandoval, 9 Nov 2008. MEPN 11111 (7, 55.2 – 124.9 mm SL): Zamora- Chinchipe, Sacha Vaca Alta, Río Machinaza, 3 ° 40 ’ 53 ’’ S, 78 ° 29 ’ 25 ’’ W; M. Andrade, 3 Aug 2011. MEPN 11113 (6, 147.3 – 191.7 mm SL): Zamora-Chinchipe, Río Machinaza, 3 ° 45 ’ 46 ’’ S, 78 ° 29 ’ 46 ’’ W; R. Barriga et al., 13 Sept 2009. MEPN 11119 (20, 148.3 – 225.0 mm SL): Zamora-Chinchipe, Rio Machinaza, Peñas; R. Barriga et al., 15 Feb 2010. MEPN 11120 (2, 158.1 – 185.9 mm SL): Zamora-Chinchipe, Río Machinaza, Sachavaca, c. 3 ° 42 ' S, 78 ° 29 ' W; M. Andrade, 29 Jul – 3 Aug 2011. MEPN 11121 (3, 132.8 – 158.5 mm SL): Zamora-Chinchipe, Río Blanco, below the bridge, San Antonio, 3 ° 50 ' 49 '' S, 78 ° 41 ' 41 '' W; R. Barriga et al., 18 July 2010. MEPN 11117 (4, 39.8 – 121.2 mm SL): Río Tundayme, trib. Río Santiago, 3 ° 35 ’ 29 ’’ S, 78 ° 28 ’ 10 ’’ W; no collector specified, 7 Aug 2010. MEPN 11118 (1, 48.0 mm SL): Río Tundayme, trib. Río Santiago, 3 ° 34 ’ 35 ’’ S, 78 ° 28 ’ 47 ’’ W; no collector specified, 6 Aug 2010. Província Morona-Santiago: MEPN 1979 (4, 119.1 – 132.9 mm SL): Morona-Santiago, Río Chuchumbletza, 3 ° 32 ’ 20 ’’ S, 78 ° 30 ’ 20 ’’ W; R. Barriga, 13 Sept 1978. MEPN uncataloged (1, 50.2 mm SL): Quebrada Chuza, 3 km from Tayuza, c. 2 ° 43 ’ S, 78 ° 15 ’ W; R. Barriga, 7 May 1991. MEPN 11117 (2, 147.9 – 177.1 mm SL): Río Chiguaza, trib. Río Paztaza, 2 ° 1 ' 30 '' S, 77 ° 59 ' 38 '' W; M. Olalla, 8 Aug 1956. MEPN 5239 (22, 27.8 – 47.2 mm SL): Rio Chupianza, 3 km from Chupianza, 2 ° 43 ’ 31 ’’ S, 78 ° 18 ’ 48 ’’ W; R. Barriga, 6 March 1991. MEPN 5190 (1, 56.0 mm SL): Río Paute at the mouth of río Napiñanza, 2 ° 39 ’ 20 ’’ S, 78 ° 23 ’ 18 ’’ W; R. Barriga, 7 May 1991. Peru, Depto. Amazonas, Provincia Condorcanqui: MUSM 6136 (10, 52.8 – 135.8 mm SL); MUSM 6153 (3, 97.3 – 117.8 mm SL); MUSM 6163 (1, 104.0 mm SL); MUSM 6144 (4, 75.0 – 108.2 mm SL); MUSM 6141 (1, 145.1 mm SL); MUSM 6123 (7, 53.4 – 103.4 mm SL): Río Comainas, tributary of Río Cenepa, at Posto de Vigilancia 22, c. 3 ° 57 ’ 40 ’’ S, 78 ° 24 ’ 40 ’’ W; H. Ortega et al., 24 – 28 Jul 1994. MUSM 6127 (8, 82.8 – 135.4 mm SL); ZUEC 6851 (2, 96.5 – 105.4 mm SL): Río Comainas, quebrada 3, Posto de Vigilancia 22, 3 ° 57 ’ 40 ’’ S, 78 ° 24 ’ 40 ’’ W; H. Ortega et al., 21 Jul 1994.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F6DFFD84EA4F9EAFA9BFC2F.taxon	diagnosis	Diagnosis. Brycon polylepis can be distinguished from all remaining cis-andean Brycon species, except B. hilarii, B. orbignyanus, and B. orthotaenia, by possessing a dark caudal peduncle stripe extending onto the middle caudalfin rays. It can be distinguished from all these species by possessing narrow longitudinal stripes along the body straight, formed by pigmentation concentrated in the medio-distal portion of scales (vs. longitudinal stripes along the body wavy, formed by pigmentantion concentrated on upper and lower portions of scales), and 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 (vs. 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). Additionaly, Brycon polylepis can be diagnosed from most cis-andean Brycon species by its high lateral line counts (68 – 94, modally 76).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F6DFFD84EA4F9EAFA9BFC2F.taxon	description	Description. Morphometric data are presented in Table 3. Middle-sized species, largest examined specimen 248.0 mm SL. Body slender. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly convex to straight from latter point to basis of supraoccipital process, and slightly to moderately convex from latter point to dorsal-fin origin, straight 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. Maxillary extending posteriorly to vertical at anterior third of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Six (5), 7 (14), 8 (9), or 9 (1) tricuspidate teeth in outer series. Three (20), 4 (5), or 5 (1) tricuspidate teeth in second, inner premaxillary row, plus 3 (4) or 4 (24) tricuspidate teeth between the first and third rows. Two teeth in third premaxillary row, medial teeth largest, tetracuspidate, symphyseal teeth smaller, tricuspidate. Maxillary margins approximately parallel, straight in profile. Ten to 18 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 5 (1), 6 (2), 7 (3), or 8 (1) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, tri- to tetracuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, tri- to unicuspidate. Inner (lingual) series consisting of a relatively large single unicuspid symphyseal tooth, approximately as large as the symphyseal dentary teeth of main series situated immediately anterior to it, plus row of 3 (1), 4 (3), 5 (2), or 6 (1) small, unicuspidate teeth, originating on lingual crest of dentary replacement trench at level of the smaller teeth of main series. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Sixty-eight (1), 69 (1), 75 (5), 76 (8), 77 (5), 78 (1), 79 (1), 80 (1), 82 (3), 85 (1), 91 (1), or 94 (1) scales in lateral line series; laterosensory tube simple, slightly deflected downwards. Horizontal scale rows between dorsal-fin origin and lateral line 11 (7), 12 (18), 13 (3), or 14 (1). Horizontal scale rows between lateral line and pelvic-fin 7 (10), 8 (15), 9 (1), or 10 (2). Circumpeduncular scales 20 (4), 21 (10), 22 (6), 23 (5), 24 (1), or 25 (2). Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead of middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 11 th vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 25 (1), 26 (2), 27 (10), 28 (11), 29 (2), 30 (2), or 31 (1). First anal-fin pterygiophore inserting behind haemal spine of 24 th vertebra. Last unbranched and anterior 3 – 4 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 10 – 30 per fin-ray main branch) small hooks on last unbranched and posterior main branch of branched rays 19 – 20, associated with dense, gelatinous tissue in two specimens (MBUCV-V 15094, 197.1 mm SL; CAS 68838, 190 mm SL). Generally a single, and at most three, hooks per ray segment. Sheath of scales composed of three scale rows covering basis of anal-fin rays, inferior scale row larger. Inferior scale row formed by 25 – 28 rectangular scales. Pectoral-fin rays i, 11 (3), 12 (21), or 13 (4). Pelvic-fin rays i, 7. Branched pelvic-fin rays with small hooks (c. 15 – 30 per branch) on distal portion of posterior ray branches in two specimens (MBUCV-V 15094, 197.1 mm SL; CAS 68838, 190.0 mm SL). Main caudal-fin rays 10 / 9. Caudal fin forked, lobes rounded to slightly pointed. Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First gill arch with 4 hypobranchial, 12 ceratobranchial, 1 on angle, 12 epibranchial, and 4 on cartilage between pharygobranchial and ceratobranchial gill rakers. Vertebrae 46 (1). Supraneurals 12 (1), middle supraneurals with a moderately developed lateral flanges. Coloration in alcohol. Top of head, snout, supraorbital, first and sixth infraorbitals, upper portion of fifth infraorbital and opercle, and dorsal portion of body chestnut- to dark-brown. Second, third, and fourth infraorbitals, lower half of fifth infraorbital, and lower opercle two-thirds silvery. Lateral portion of body clear, with silvery and brownish hues. Dentary, maxillary, gular area, and lower portion of body clear, with a brownish hue. Dark chromatophores concentrated at lower portion of fifth infraorbital bone and middle portion of opercle forming a relatively broad, faint postorbital dark stripe in some specimens. Humeral blotch present, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of third, extending longitudinally to posterior margin of fifth to sixth lateral line scales, and vertically one and a half scales high. Specimens around 200 mm SL (e. g., MZUSP 59000, MZUSP 88325) possess longitudinal, narrow straight stripes across the body formed by pigmentation concentrated on the central-distal portion of the scales. Longitudinal stripes less conspicuous in smaller specimens. Oblique, narrow vertical stripes resulting from coalescence of dark pigmentation concentrated on the central-distal portion of scales present in some specimens (e. g., MZUSP 88325, MZUSP 56784, MBUCV-V 15094). Broad, conspicuous caudal peduncle dark stripe, approximately rectangular in profile, extending to distal portion of 5 – 6 medialmost caudal-fin rays. Adipose fin cream colored. Remaining fins translucent, except for dark pigmentation moderately developed in anal-fin interradial membranes in some specimens (e. g., MUSM 11077; MUSM 33345). Specimens retaining most of the original guanine pigmentation (e. g., MUSM 33345) silvery overall, with dorsal portion plumbeous-grey. Color in life. Based on a picture of specimen (not preserved) fished at the Río Negro, a tributary of Río Casanare, Río Orinoco basin, Colombia, provided by A. Linares, and a specimen from the Río Cataniapo, upper Río Orinoco basin, depicted by Fernández et al. (2006: 57). Overall body coloration clear, with a strong silvery hue. Tof of head, snout and dorsal region grey. Upper portion of opercle with a yellow blotch. All fins pinkish, with some yellow pigmentation at their basis. Sexual dimorphism. Two examined specimens (MBUCV-V 15094, 197.1 mm SL and CAS 68838, 190.0 mm SL) were recorded displaying fin hooks on the anal and pelvic-fins. None of these specimens was dissected to verify their sex, but Moscó Morales (1988: 20), who examined several additional specimens that were not examined in the present study, confirmed that the males possess, while the females do not, hooks in the anal- and pelvic-fins. Variation. Two specimens collected at the Río Nanay, a black-water tributary of the Amazon River near Iquitos (INHS 44001) are distinct from the remaining examined specimens by possessing an almost straight dorsal profile (vs. slightly convex profile), a lower body depth (20.2 – 20.4 % SL, vs. 23.7 – 29.2 % SL), and a slightly distinctly shaped caudal-peduncle stripe (slightly oblique, extending into lower portion of caudal peduncle and onto its lower surface, vs. approximately straight stripe, not extending into lower portion of caudal peduncle and onto its lower surface). Additionally, these specimens possess an average high number of lateral-line scales (85 – 91, vs. 68 – 82, median 76 scales for the remaining examined specimens). However, a specimen from the upper Río Marañon in Peru (MUSM 33345) possess a lateral-line scale count even higher (94 scales) than the specimens from the Río Nanay, without sharing with them a low body depth or a straight dorsal profile. More specimens of this scarce species are necessary for a better understanding of its intraspecific variation. Common names. Brazil, Goiás: “ piabanha ” (pers. obs.); Venezuela, Estado Zulia: “ palambra ”, “ dientón ” (Moscó Morales, 1988: 9); Estado Amazonas: “ bocón-rey ”, “ palambra ”, “ Icuä äi ” (Piaroa language) (Fernández et al., 2006: 57).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F6DFFD84EA4F9EAFA9BFC2F.taxon	distribution	Distribution. Described originally from Río Santa Ana and Río Limón basins, Lago de Maracaibo drainage, Venezuela, and known from scattered localities in the Río Orinoco basin in Venezuela and Colombia (record based on a photograph of a specimen angled at the upper Río Negro, a tributary of Rio Casanare, itself a tributary of Rio Meta; aproximately 6 ° 4 ’ N, 72 ° 9 ’ W; A. Linares, pers. comm.), rio Tocantins basin in Brazil, and the upper Amazon basin in tributaries of Rio Madeira, Maranõn and Amazonas in Peru (Fig. 20). The species is very probably widely distributed in piedmont localities across the upper Río Orinoco and Rio Amazonas basins. Ecological notes. Most known specimens of Brycon polylepis were collected in headwater streams and small rivers possessing clear water, swift current and generally a rocky bottom (Moscó Morales, 1988: 20; Fernández et al., 2006: 57; pers. obs.). Specimens MZUSP 18142 were collected in a rocky pool at the rapids of Jatobal, a former extensive rapid area of the rio Tocantins now submerged under the Tucuruí dam. Curiously, as reported by Bichuette & Trajano (2003; as Brycon sp.), a small population of the species was recorded as living for several years in a cave stream at the São Domingos karst area in the upper rio Tocantins basin in Brazil; these specimens were recorded feeding cave crickets accidentally fallen in the water (pers. obs.; E. Trajano, pers. comm ..). Streams in the karst zone of São Domingos run across the limestone outcrops of the Bambuí formation, heading towards the rio Paranã, one of the upper tributaries of the rio Tocantins (see Bichuette & Trajano, 2003, fig. 1), and Brycon polylepis was recorded from both sinkhole and resurgence areas in these streams (specimens MZUSP 59000, MZUSP 88325, and MZUSP 88326). Moscó-Morales (1988: 20) reported leaves, trichopteran cases, fruits, seeds, and insect remains in stomach contents he examined. Fernández et al. (2006: 57) reported, without further details, that the reproduction of the species at the Río Cataniapo had it onset during the beginning of the rainy season.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F6DFFD84EA4F9EAFA9BFC2F.taxon	discussion	Remarks. Comparisons between the single examined paratype of Brycon polylepis from the Lake Maracaibo basin and specimens from the upper Amazon basin in Peru, rio Tocantins basin in Brazil, and Río Orinoco basin in Venezuela did not revealed any distinguishing characters between the respectively trans- and cis-andean populations of Brycon polylepis, and these specimens are accordingly referred to a single species. See the section “ Biogeography ”, below.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F6DFFD84EA4F9EAFA9BFC2F.taxon	materials_examined	Material examined. Type material: MBUCV-V 15094 (1 of 2, 197.1 mm SL): Venezuela, Estado Zulia, Río Yasa, tributary of Río Negro (tributary of Rio Santa Ana), Estacion Biologica de Kasmera, Sierra de Perijá, c. 9 ° 57 ’ N, 72 ° 43 ’ W; J. Moscó Morales et al., 20 Sept 1985. Paratype of Brycon polylepis Moscó Morales, 1988. Non types: Brasil, rio Tocantins basin, Pará: MZUSP 56784 (1, 132.2 mm SL): Parauapebas, igarapé Salobo (tributary of rio Itacaiúnas), 5 ° 46 ' 45 '' S, 50 ° 32 ' 13 '' W; P. S. Pompeu, Nov 1997. ZUEC 7065 (1, 121.7 mm SL): Parauapebas, igarapé Salobo (trib. rio Itacaiúnas), 5 ° 49 ’ 33 ’’ S, 50 ° 29 ’ 25 ’’ W; T. Giarizzo, 15 Dec 2009. ZUEC 7064 (1, 105.4 mm SL): Parauapebas, igarapé Mirim (trib. rio Itacaiúnas), 5 ° 45 ’ 36 ’’ S, 50 ° 30 ’ 59 ’’ W; T. Giarizzo, 15 Sept 2009. MZUSP 18142 (1, 106.9 mm SL; 1, 104.4 mm SL, cs), rio Tocantins, lagoon at Jatobal, 4 ° 32 ’ S, 49 ° 28 ’ W; EPA, 16 Sept 1970. INPA 16391 (1, 100.6 mm SL), igarapé Valentim, tributary of rio Tocantins, Tucuruí-Marabá road, km 130; G. M. Santos, 6 July 1982. Goiás: MCP 16008 (3, 113.8 – 122.4 mm SL), Niquelândia, Ribeirão Arara, 500 m from its mouth in the rio Maranhão at Rosariana; 14 ° 01 ' S 48 ° 25 ' W; R. E. Reis et al., 14 July 1992. MZUSP 59000 (1, 202.7 mm SL), São Domingos, entrance of Lapa da Bezerra, Terra Ronca State Park, 13 ° 32 ' 43 '' S, 46 ° 22 ' 35 '' W; A. Akama, 24 June 1999. MZUSP 88325 (1, 189.4 mm SL), São Domingos, Rio da Lapa at its resurgence, Caverna Terra Ronca, Terra Ronca State Park; M. E. Bichuette & R. Santos, 14 Sept 1999. MZUSP 88326 (2, 200.9 – 201.3 mm SL), São Domingos, Rio da Lapa, at entrance of Caverna Terra Ronca, Terra Ronca State Park, 13 ° 44 ’ 00 ’ S, 46 ° 21 ’ 30 ’’ W; M. E. Bichuette & R. Santos, 14 Sept 1999. MNRJ 13069 (4, 109.2 – 125.2 mm SL), Formosa, rio Água Quente (tributary of rio Paranã), at the crossing of the Formosa-Itiquira- Flores de Goiás road, 74 km from Formosa, 14 ° 58 ' 23 '' S, 47 ° 28 ' 36 '' W; L. E. M. Cardoso, 4 Sept 1982. IBGE uncat. (1, 107.0 mm SL): São João da Aliança, stream trib. rio Paranã, fazenda Farias, 14 ° 31 ' 18 ' ’ S, 47 ° 10 ' 36 ' ’ W; M. C. L. Ribeiro et al., 14 June 1990. MZUSP 70413 (2, 192.2 – 203.9 mm SL), rio Tocantins, downstream UHE Serra da Mesa, 13 ° 49 ' 54.8 '' S, 48 ° 17 ' 44.2 '' W; D. F. Moraes & D. A. Halboth, 4 – 5 Feb 1997. MZUSP 70414 (1, 120.6 mm SL), rio Tocantizinho, Serra da Mesa dam, 13 ° 57 ' 10 '' S, 48 ° 19 ' 33 '' W; D. F. Moraes & D. A. Halboth, 4 Dec 1997. MZUSP 70419 (1, 103.7 mm SL), rio Maranhão, near rio do Peixe mouth, 14 ° 14 ' 54.9 '' S, 48 ° 55 ' 39.7 '' W; D. F. Moraes & F. P. Matos, 12 Feb 1997. MZUSP 71671 (1, 90.2 mm SL), córrego do Cavalo, tributary of rio Tocantins, above Serra da Mesa dam, 14 ° 31 ' 16 ' ’ S, 48 ° 56 ' 6 ' ’ W; J. C. Miranda et al., 11 Feb 1997. IBGE uncat. (1, 200.0 mm SL): Padre Bernardo, Rio do Sal, trib. rio Maranhão, 15 ° 20 ’ 20 ’’ S, 48 ° 3 ’ 5 ’’ W; M. C. L. Ribeiro et al., 6 Aug 1998. CAS 68838 (1, 190 mm SL); CAS 68834 (1, 200 mm SL): upper Rio Maranhão at “ Mosondo ” (not located), into rio Tocantins; C. Ternetz, 2 – 4 Oct 1923. CAS 68823 (1, 208 mm SL), " Tirap-lanya " (not located), rio Maranhão, into rio Tocantins; C. Ternetz, 2 Oct 1923.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F6DFFD84EA4F9EAFA9BFC2F.taxon	description	Peru: INHS 44001 (2, 109.7 – 124.1 mm SL), Depto. Loreto, Río Nanay, Pampa Chica, 4.54 km W center of Iquitos, 3 ° 45 ' 08.8 '' S 73 ° 17 ' 00.1 '' W; M. H. Sabaj & J. W. Armbruster, 22 July 1997. MUSM 11077 (1, 206.0 mm SL): Depto. Puno, Sandi, Zona Reservada Tambopata / Candamo, Río Candamo (tributary of Rio Tambopata), quebrada Ebebahuaeji, c. 13 ° 24 ' S, 69 ° 35 ' W; F. Chang, 31 March 1997. MUSM 33345 (1, 248.0 mm SL): Depto. San Martin, Prov. M. Cáceres, Huicungo, Río Abiseo (tributary of Rio Huallaga), Río Abiseo National Park, 7 ° 26 ' 56 '' S 76 ° 53 ' 43 '' W; H. Ortega, A. Díaz & P. Zuñiga, 20 May 2008. MUSM 43197 (1, 123.3 mm SL): Depto. Ucayali, Coronel Portillo, Río Shesha (trib. Río Ucayali), 8 ° 11 ’ 17 ’’ S, 73 ° 56 ’ 38 ’’ W; V. Meza et al., 17 Oct 2011. Venezuela, Río Orinoco basin: ANSP 159717 (1, 93.7 mm SL), Bolivar, caño 15.1 km E of Río Parguaza, ferry crossing on Caicara-Puerto Ayacucho hwy, 6 ° 26 ’ 28 ’’ N, 67 ° 9 ’ 24 ’’ W; B. Chernoff et al., 28 Nov 1985.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F57FFDB4EA4FBDBFD29FF10.taxon	diagnosis	Diagnosis. Brycon coquenani can be distinguished from all remaining cis-andean Brycon species, except B. stolzmanni, B. coxeyi, B. devillei, B. insignis, B. vermelha, Brycon howesi, B. dulcis, B. ferox, Brycon vonoi, B. opalinus, and B. nattereri, by possessing a color pattern composed by a humeral blotch and a caudal peduncle blotch (vs. body stripes and caudal / anal fin color markings present; see Fig. 5). Brycon coquenani can be distinguished from all these species, except B. insignis, B. vermelha, and B. howesi, by possessing a fifth infraorbital bone wider than high (vs. fifth infraorbital bone about as wide as high; see Fig. 6). Brycon coquenani can be distinguished from these species by possessing pointed, tri- to tetracuspidate dentary teeth, with lateral cusps poorly developed (vs. pointed tri- to pentacuspidate dentary teeth, with moderately developed lateral cusps in B. insignis, B. vermelha, and B. howesi), and by mature males displaying hooks in all rayed fins (vs. hooks present generally only in the anal-fin, rarely also in the pelvic-fins in mature males in B. insignis, and B. howesi, and not present in B. vermelha).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F57FFDB4EA4FBDBFD29FF10.taxon	description	Description. Morphometric data are presented in Table 4. Relatively small-sized species, largest examined specimen 170.6 mm SL. Body moderately slender. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly concave from latter point to basis of supraoccipital process, and moderately convex from latter point to dorsal-fin origin, straight along dorsal-fin basis, straight to slightly convex from dorsal-fin terminus 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. Maxillary extending posteriorly to vertical slightly anterior to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Small specimens (<84.2 mm SL) with gaps (diastemas) between teeth. Eight (1), 9 (5), 10 (2), or 11 (1) unicuspidate teeth in outer series. Three (2), 4 (2), or 5 (1) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 1 (1), 2 (3), or 3 (1) tricuspidate teeth between the first and third rows. Two teeth in third premaxillary row, medial teeth largest, pentacuspidate, symphyseal teeth smaller, tricuspidate. Maxillary with distal portion expanded and rounded in profile. Twenty-five to 31 maxillary teeth, unicuspidate, slightly smaller than teeth of first premaxillary row. Dentary with 8 (1), 9 (2), 10 (1), 11 (3), or 12 (2) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger than remaining teeth, tri- to tetracuspidate. Remaining dentary teeth progressivelly smaller, unicuspidate. Inner (lingual) series consisting of a single unicuspid symphyseal tooth, considerably smaller than teeth of main series anterior to it, plus a row of small, unicuspidate teeth. All teeth with well-developed, pointed central cusp, remaining cusps, when present, poorly developed or vestigial. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-four (1), 46 (3), 47 (2), or 48 (3) scales in lateral line series; laterosensory tube simple, straight or deflected downwards. Horizontal scale rows between dorsal-fin origin and lateral line 8 (6), or 9 (3). Horizontal scale rows between lateral line and pelvicfin 4. Circumpeduncular scales 17 (2), or 18 (1). Dorsal-fin rays ii, 8 (2) or 9 (9). Dorsal fin origin slightly ahead from middle of SL. Dorsal fin displaying numerous (c. 15 – 30 per fin-ray main branch) small hooks on last unbranched and posterior main branch of all branched rays in one specimen (lectotype, NMW 62702, 170.6 mm SL). Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 18 (1), 19 (2), 21 (3), 22 (4), 23 (2), or 24 (2). Last unbranched and anterior 2 – 3 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 20 – 30 per fin-ray main branch) small hooks associated with dense, gelatinous tissue on last unbranched and posterior main branch branched rays to antepenultimate branched ray in one specimen (lectotype, NMW 62702, 170.6 mm SL). Sheath of scales composed of a single series composed by 15 rectangular scales, covering basis of anal-fin rays. Pectoral-fin rays i, 11 (2), 12 (4), or 13 (3). Branched pectoral-fin rays with minute hooks (c. 10 per branch) on distal portion of posterior ray branches in one specimen (lectotype, NMW 62702, 170.6 mm SL). Pelvic-fin rays i, 7. Branched pelvic-fin rays with minute hooks (c. 10 – 20 per branch) on distal portion of posterior ray branches in one specimen (lectotype, NMW 62702, 170.6 mm SL). Main caudal-fin rays 10 / 9. Small hooks (c. 5 – 10 per ray branch) on distal portion of inner caudal-fin rays in one specimen (lectotype, NMW 62702, 170.6 mm SL). Caudal fin forked, lobes rounded. Coloration in alcohol. Color description based on the lectotype, NMW 62702; paralectotypes too faded to allow an accurate color description. Top of head, snout, supraorbital, sixth infraorbital bone, and dorsal portion of body dark brown. Dentary, maxillary, first, second, and upper portion of fifth infraorbital bone, gular area, and ventral portion of body cream-colored. Third, fourth, lower portion fifth infraorbital bones, and opercle silvery, with a brownish tinge. Humeral blotch present, relatively inconspicuous, rounded, extending from first through third lateral line scales. Lateral portions of body cream-colored, with a silvery hue. Caudal peduncle blotch present, conspicuous, rounded, extending throughout 6 last lateral line scales into basis of 4 innermost caudal-fin rays. Caudal, anal and dorsal-fins with numerous small dark chromatophores scattered over interradial membranes. Pectoral and pelvic-fins with relatively few dark chromatophores on interradial membranes. Adipose fin light brown. Common name. “ Aruma ”. This common name, mentioned by Steindachner (1917: 37), was recorded by the collector of the type material, John Haseman. In an undated manuscript authored by Haseman, entitled “ Zoogeographical expedition to northeastern South America ”, he mentioned (p. 44) “ aruma ” as the “ native ” name of an undescribed species of Brycon collected in the Río Kukenan. The Amerindians which helped Haseman during his expedition into the tepuis of northern Roraima state in Brazil and southern Bolívar state in Venezuela belonged to the Arekuna (= Taulipang), so it can be assumed that this common name is originated from this language. Sexual dimorphism. The lectotype (NMW 62702, 170.6 mm SL) possess numerous small hooks over all rayed fins, including the caudal-fin. However, a dissection was not performed to confirm the specimen’s sex.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F57FFDB4EA4FBDBFD29FF10.taxon	distribution	Distribution. Brycon coquenani is only known from the upper Rio Kukenan (“ Coquenanflusse ”), a tributary of the upper Río Caroní, Río Orinoco basin, estado Bolívar, Venezuela (Fig. 20). The exact locality of Brycon coquenani is not known but according to the narrative of Haseman’s travel in the region referred above, the type series was collected in the Rio Kukenan at its valley between the mount Roraima and the Cerro Kukenan, below the “ Moromelu Falls ” (not located).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F57FFDB4EA4FBDBFD29FF10.taxon	discussion	Remarks. Steindachner (1917: 37 – 38) described the species based on a syntypical series of “ 11 ... juveniles and a larger, very well preserved exemplar 210 mm long ” (our translation). We have examined the syntypical series of Brycon coquenani at the NMW. The largest specimen, NMW 62702, is herein designated as the lectotype of the species. The remaining, smaller syntypes, NMW 62703 (10 specimens) became, consequently, paralectotypes. The lectotype is in very good condition, except for missing some scales and for having the top of head bruised, apparently by Steindachner himself, presumably for the examination of the fontanel. In spite of relatively recent collecting efforts in the upper Río Caroní basin, including the Río Kukenan, so far no additional specimens of the species were obtained (Lasso, 1990; Lasso et al., 1990).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F57FFDB4EA4FBDBFD29FF10.taxon	materials_examined	Material examined. Type material: NMW 62702 (1, 170.6 mm SL): Venezuela, Estado Bolívar, Río Kukenan, upper Río Caroní basin, Gran Sabana, approximately 5 ° 12 ’ N, 60 ° 46 ’ W; J. D. Haseman, April 1913. Lectotype of Brycon coquenani Steindachner, by present designation. NMW 62703 (10, 64.7 – 84.2 mm SL): same data as lectotype. Paralectotypes of Brycon coquenani.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F54FFD24EA4FE8BFC07FCF3.taxon	description	Megalobrycon piabanga (sic): Magalhães, 1931: 155 – 156, fig. 81 (Rio Paraíba do Sul; natural history).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F54FFD24EA4FE8BFC07FCF3.taxon	description	Brycon devillei (non Castelnau): Araújo, 1996: 116, 119, 121 (Rio de Janeiro, rio Paraíba do Sul, São Fidélis).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F54FFD24EA4FE8BFC07FCF3.taxon	diagnosis	Diagnosis. Brycon insignis can be diagnosed from all remaining cis-andean Brycon species, with the exception of B. stolzmanni, B. coxeyi, B. coquenani, B. vermelha, B. howesi, B. dulcis, B. ferox, B. vonoi, B. opalinus, and B. nattereri by possessing a color pattern consisting in a humeral blotch and a caudal peduncle blotch, without body stripes or other obvious color markings on caudal and anal-fins (vs. body stripes and caudal / anal fin color markings present) (see Fig. 5). Brycon insignis can be diagnosed from these species, with the exception of B. howesi, B. vermelha, and B. coquenani, by possessing a fifth infraorbital bone considerably wider than high (vs. fifth infraorbital bone about as wide as high; see Fig. 6). Brycon insignis can be diagnosed from B. coquenani by possessing a higher head (63.1 – 75.9 % of head length, mean 68.9, vs. 59.8 – 64.9 %, mean 61.4, in B. coquenani), by possessing pointed tri- to pentacuspidate dentary teeth, with moderately developed lateral cusps (vs. pointed, trito tetracuspidate dentary teeth, with lateral cusps poorly developed; see, however, the item “ Remarks ”, below, on individuals formely assigned to Catabasis acuminatus), and by mature males displaying hooks only in anal fin (vs. mature males displaying hooks in all rayed fins, including caudal fin). Brycon insignis can be distinguished from B. vermelha by presenting darkened dorsal, caudal, and anal-fins in both living and preserved specimens (vs. dorsal, caudal, and anal-fins reddish in living, clear in preserved, B. vermelha specimens). Brycon insignis can be distiguished from Brycon howesi by possessing 51 – 65, modally 58 lateral line scales (vs. 46 – 50, modally 49 in B. howesi), 9 – 11, modally 10 horizontal scale rows between the dorsal-fin origin and the lateral line (vs. 7 – 9, modally 8 in B. howesi), 4 – 7, modally 5 horizontal scales rows between the lateral line and pelvic fin (vs. 3 – 5, modally 4 in B. howesi) and a pointed snout in profile in B. insignis (vs. snout pointed, but rounder, in B. howesi). For more comparisons among Brycon insignis, B. vermelha, and B. howesi, see the item “ Remarks ” of these latter two species.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F54FFD24EA4FE8BFC07FCF3.taxon	description	Description. Morphometric data are presented in Table 5. Large-sized species, largest examined specimen 493.0 mm SL. Body moderately slender. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, moderately convex from latter point to basis of supraoccipital process, and 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 conspicuously acute anteriorly, mouth terminal. Jaws anisognathous, premaxillary projecting slightly relative to dentary, outer row of premaxillary teeth and in some specimens also second row exposed when mouth is closed. Maxillary long, extending posteriorly to slightly posterior to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Five (1), 7 (6), 8 (12), 9 (13), 10 (9), 11 (6) ou 12 (3) tricuspidate teeth in outer series. Three (1), 4 (21), 5 (18), 6 (6), or 7 (1) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 1 (14), 2 (23), or 3 (12) 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, both pentacuspidate. Maxillary with distal portion distinctly expanded and rounded in profile. Eighteen to 33 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 9 (1), 10 (1), 11 (4), 12 (7), 13 (5), 14 (10), 15 (10), 16 (6), or 17 (3) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, tetra- to pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, tetra, tri- 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 23 – 37 small, unicuspidate teeth, originating on lingual crest of dentary replacement trench at level of the fourth to sixth teeth of main series. Inner symphyseal teeth absent in 6 out of 19 specimens examined for this character. Larger specimen displaying symphyseal teeth 220.3 mm SL (MZUSP 2091); specimens larger than 230 mm SL (MCZ 21112, MZUSP 103029, MZUSP 88515, MZUSP 19128) lacking symphyseal teeth. A single specimen smaller than 220 mm SL lacking symphyseal teeth (MNRJ 4716, 138.3 mm SL). Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Fifty-one (2), 52 (2), 53 (5), 54 (6), 55 (5), 56 (8), 57 (5), 58 (9), 59 (5), 60 (2), 61 (2), 62 (2), 63 (4) ou 65 (1) scales in lateral line series. Laterosensory tube simple in small (<200 mm SL) specimens, deflected upwards in the first 5 – 6 scales, downwards in the remaining lateral-line scales. Larger specimens (> 200 mm SL) with branched tubules, mostly bifurcated in middle-sized (230 – 270 mm SL) specimens but some scales with 3 – 5 tubules in the larger (> 290 mm SL) specimens examined. Horizontal scale rows between dorsal-fin origin and lateral line 9 (9), 10 (25), or 11 (22). Horizontal scale rows between lateral line and pelvic-fin 4 (7), 5 (27), 6 (17), or 7 (3). Circumpeduncular scales 16 (3), 17 (2), 18 (9), 19 (10), 20 (12), 21 (12), or 22 (3). Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead of middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 13 th vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 21 (2), 22 (2), 23 (10), 24 (9), 25 (12), 26 (17), or 27 (5). First anal-fin pterygiophore inserting behind haemal spine of 24 th vertebra. Last unbranched and anterior 3 – 4 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 18 – 22 per fin-ray main branch) middle-sized hooks on last unbranched and posterior main branch of branched rays 10 – 20, associated with dense, gelatinous tissue in six specimens (MZUSP 2091, 209.0 – 220.3 mm SL; MZUSP 79092, 187.2 mm SL; MZUSP 88515, 235.0 – 247.0 mm SL; MZUSP 107280, 260.0 mm SL). A single hook per ray segment. Sheath of scales covering basis of anal-fin rays composed of two scale rows, lower scale row formed by 25 – 30 rectangular scales. Pectoral-fin rays i, 11 (1), 12 (10), 13 (27), 14 (15), or 15 (1). Pelvic-fin rays i, 7. Main caudal-fin rays 10 / 9. Caudal fin forked, lobes slightly pointed. Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First branchial arch with 11 (2), 12 (6), 13 (10), 14 (12), 15 (3), or 16 (1) lower, 1 at angle, and 11 (4), 12 (14), 13 (12), 14 (3), or 17 (1) upper gill rakers. Vertebrae 45 (2). Supraneurals 9 (1). Coloration in alcohol. Description based on wild-caught, recently collected specimens (MZUSP 88515 and MZUSP 103045); the majority of remaining examined specimens faded, either due to a long period of conservation (> 60 years) or for being originally captive, stocked specimens raised in clear-water, sun-lit ponds. Top of head, snout, supraorbital, sixth infraorbital, upper portion of fifth infraorbital, and dorsal portion of body dark-brown. Second, third, and fourth infraorbitals light brown, with a silvery hue. Opercle mainly silvery, with some lightbrown tinge. Dentary, maxillary, gular area and lower portion of body clear, with a cream tinge. Lateral portion of body light-grey, silvery in specimens retaining a considerable amount of guanine (e. g., MCZ 21112, MZUSP 62746). Scales above lateral line on lateral surfaces of body with dark pigmentation concentrated on scales margins, forming a tenuous reticulated pattern in specimens MZUSP 88515 and MZUSP 103045. Humeral blotch present, moderately 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 fourth to fifth lateral line scales, and vertically less than one scale high. Lateral surfaces of body above lateral line, and posterior to dorsalfin, presenting longitudinal stripes formed by dark pigmentation on central portion of scales in specimens MZUSP 88515 and MZUSP 103045. Large, moderately conspicuous, oval-shaped caudal peduncle blotch, extending along 8 – 11 last lateral-line scales. All rayed fins with a considerable amount of dark pigmentation at the interradial membranes (discernible even in some severely faded specimens, e. g., MZUSP 2091), imparting a overall dark coloration to these fins. Caudal-fin with a poorly discernible, roughly V-shaped blotch, formed by dark pigmentantion situated on outer caudal-fin rays. Adipose fin light-grey to light-brown. Color in life. Description based on a specimen photographed alive at the CESP-Paraibuna fish hatchery and from photographs of specimens MZUSP 103045, immediately before fixation. Top of head and snout dark. Dentary and maxillary clear. Dorsal portion of body plumbeous. Opercle, infraorbital bones, and lateral portion of body silvery. Humeral blotch inconspicuous; caudal peduncle blotch moderately conspicuous. All fins, except adipose fin, darkened. Caudal fin with outer fin-rays dark, forming a roughly V-shaped blotch. Sexual dimorphism. Five specimens (MZUSP 2091, 209.0 – 220.3 mm SL; MZUSP 79092, 187.2 mm SL; MZUSP 88515, 235.0 – 247.0 mm SL; MZUSP 107280, 260.0 mm SL) possess anal-fin hooks. Hooks are, however, absent from pelvic- and remaining fins. One specimen (MZUSP 2091) proved to be a male after dissection. At the CESP-Paraibuna fish hatchery, mature males were in fact distinguished from females due to the presence of analfin hooks (L. Girardi, pers. comm, 1999). Common names. “ Piabanha ”; “ piabanha-vermelha ” (Miranda-Ribeiro, 1902: 254); “ biririca ” (juvenile specimens, rio Itabapoana, V. L. M. Santos, pers. comm.).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F54FFD24EA4FE8BFC07FCF3.taxon	distribution	Distribution. Known from the rio Paraíba do Sul basin at São Paulo, Rio de Janeiro, and Minas Gerais states, plus several smaller coastal river drainages at Rio de Janeiro state, i. e., rio Guandu, rio Macaé, rio São João, rio Imbé and Lagoa Feia system, and rio Itabapoana, at the border of Rio de Janeiro and Espírito Santos states (Fig. 25). Specimens from the rio Macaé were not examined in the present study, but the species also occurs at this small river system according to Bizerril & Primo (2001). Extirpated from much of its original range (see “ Conservation ”, below). Ecological notes. Stomach contents of a dissected specimen (MZUSP 2091) contained a whole, nonmasticated fruit and flowers remains. Bizerril & Primo (2001: 55, as B. opalinus) reported insects from stomachs of specimens collected at the rio Paraíba do Sul. Magalhães (1931: 156) reported smaller fishes as the main dietary item of B. insignis. Males achieve maturity at the second year of age, with about 20 cm TL, while females mature at the third year of age, at the size of 25 cm TL (Salgado et al., 1997). Microsatellite anaylsis of populations across the rio Paraíba do Sul basin showed significant differentiation among distinct populations (Matsumoto & Hilsdorf, 2008).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F54FFD24EA4FE8BFC07FCF3.taxon	description	Conservation. Brycon insignis was reported as being a common fish in the rio Paraíba do Sul basin in the past (Magalhães, 1931). Machado & Abreu (1952) recorded that Brycon insignis was the second more important fish in the commercial fisheries in the rio Paraíba do Sul basin, with annual landings of 15 – 22 tonnes in the São Paulo portion of the basin alone. The steep decline of the populations of the species, already foreseeing by Magalhães (1931), was due to combination of several factors, the main being water pollution (both sewage and industrial pollution) and deforestation of the riparian forest. The introduction of the dourado, Salminus brasiliensis, into the rio Paraíba do Sul basin in the mid- 1950 ’ s decade is also mentioned by older fishermen as an important cause of decline of the species, probably due to competition and / or predation. The last record of Brycon insignis into the portion of the rio Paraíba do Sul basin in São Paulo state was during the late 1980 ’ s decade (Salgado et al., 1997). The species is still reported as occurring in the middle / lower stretches of the rio Paraíba do Sul basin at the Rio de Janeiro state (e. g., Bizerril & Primo, 2001; G. Sousa per A. W. Hilsdorf, pers. comm.), but some recent catastrophic fish killings due to the leakage of pesticides and other pollutants have occurred in the basin, the latter in November 2008. Brycon insignis was apparently extirpated from two small coastal river systems where it once occurred, the rio Guandu and the rio Macaé. The record for the rio Guandu is based on a single specimen collected at the former Imperial farm at Santa Cruz, during the Thayer Expedition, in 1865 (MCZ 21112). This specimen was labelled as being collected at the rio Grande, which Higuchi (1992) identified as being a small river system draining into the Lago da Tijuca, currently within the urban area of Rio de Janeiro city (Bizerril & Primo, 2001: 135 – 136). However, since the latter river lies very far from the Santa Cruz Imperial farm (more than 30 km in a straight line), we believe that the specimen was actually collected in the rio Guandu, a larger river system much closer to the imperial farm and a known locality for other Thayer Expedition specimes (as “ Rio Quenda ” or “ Quendu ”). The rio Guandu has been severely altered during the XX century and a continual occurrence of the species in this river system is highly unlikely. As for the rio Macaé, recent collecting efforts in the basin failed in locating the species (M. G. Brito, pers. comm.). Brycon insignis is officially considered as a threatened species in Brazil, under the category “ critically endangered ” (Hilsdorf, Lima & Matsumoto, 2008). Stocked populations of the species are kept in two fish hatcheries stations at São Paulo and Rio de Janeiro states (CESP-Paraibuna, Paraibuna, and Projeto Piabanha, Itaocara, respectively). There is still a reasonable genetic variability as shown by microsatellite loci in wild Brycon insignis, but considerably little variation in the captive stock of the species (Matsumoto & Hilsdorf, 2009).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F54FFD24EA4FE8BFC07FCF3.taxon	discussion	Remarks. Steindachner (1877) originally described Brycon insignis based on an unspecified number of syntypes from the “ Rio Parahyba bei Campos und Mendez, Rio Jequitinhonha ”. There are some discussion in the literature whether syntypes of Steindachner’s species described based on material collected by the Thayer Expedition should be considered as only the specimens present at the NMW collection or should also encompass material deposited at MCZ, which were examined by Steindachner during his stay at Harvard in several periods between April 1870 until his final departure from the United States in December 1873 (Steindachner, 1875 a: 499; Herzig-Straschil, 1997). Schaefer (1997: 49) considered that only specimens deposited at the NMW should be considered types, an opinion that has not been unanimously accepted. For example, Vari (1992: 48) designated a MCZ specimen as the lectotype of Steindachnerina bimaculata, quoting Steindachner’s description of the species where he refers to “ numerous specimens … in the Museum at Cambridge ” (Steindachner, 1876: 79). There are in fact a few more more references by Steindachner to material deposited at the MCZ, which points out that he has not necessarily relied exclusively on material brought by him to the NMW in his descriptions of species based totally or partially on material collected by the Thayer Expedition (for example, Leporinus mormyrops; Steindachner, 1875 b: 244). A possible compromise solution to this problem would be to recognize, as Vari (1992) did, the MCZ material as typical only in the relatively few descriptions of fishes based on material collected by the Thayer Expedition where Steindachner specifically mentioned specimens from the “ Museum zu Cambridge ”. With this view in mind, we opted for not considering as belonging to the typical series the specimens deposited at the MCZ belonging to the same original lots from which the material used by Steindachner (1877) in his descriptions of the two Brycon species based on material collected by the Thayer Expedition, B. insignis and B. ferox. We found three lots at the NMW collection which can be assigned as belonging to the syntypical series used by Steindachner (1877): NMW 62925 (2), NMW 62926 (1, 113.5 mm SL), and NMW 62927 (1, 71.6 mm SL). Label information for the lots NMW 62925 and NMW 62926 states only that they were collected at the “ Rio Parahyba ”, with no further locality information. The specimen NMW 62927 was collected at the “ Rio Jequitinhonha ”, and actually belong to the species herein described as Brycon howesi (see “ Remarks ” of this species). In order to dispel any doubts concerning the identity of Brycon insignis, we therefore select the smaller specimen of the lot NMW 62925 (144.8 mm SL), which is the syntype in better overall condition, as the lectotype of Brycon insignis, the remaining syntypes becaming, thus, paralectotypes. As noticed by Steindachner (1879 c: 50, footnote), the indication of the plate in Steindachner (1877: 135) is incorrect, Brycon insignis being actually the species portrayed in figures 1 and 1 a, figures 2 and 2 b referring to B. ferox. Steindachner (1879 c: 50) considered Brycon insignis as a synonym of B. devillei, an opinion that prevailed in the literature for a long time (e. g., Howes, 1982). Brycon devillei is considered herein as a species inquirenda (see the item “ Remarks ” under this species, below).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F54FFD24EA4FE8BFC07FCF3.taxon	materials_examined	Material examined. Type material: NMW 62925: 2 (1, 144.8 mm SL): " Rio Parahyba, Jan. 1874; Steindachner don. " Lectotype of Brycon insignis Steindachner, by present designation. NMW 62925: 1 (1, 167.7 mm SL); NMW 62926 (1, 113.5 mm SL): same data as lectotype. Paralectotypes. CAS 11894 (1, 135.1 mm SL), Brazil, São Paulo, rio Tietê; H. von Ihering, 1901 – 1905. Holotype of Catabasis acuminatus Eigenmann & Norris. Non types: Brazil, São Paulo: MZUSP 51309 (1, 253.7 mm SL); MZUSP 62746 (2, 217.9 – 310.4 mm SL); MZUSP 42045 (4, 1 cs, 143.1 – 195.8 mm SL); MZUSP 104939 (1, 493.0 mm SL): CESP-Paraibuna fish hatchery (stocked from specimens collected at Caçapava, rio Paraíba do Sul, c. 23 ° 6 ’ S, 45 ° 43 ’ W). MZUSP 19128 (1, 365.5 mm SL), Rio Paraíba do Sul, Cachoeira Paulista, c. 22 ° 39 ’ S, 45 ° 00 ’ W; J. T. Braga, May 1981. MZUSP 1553 (1, 143.1 mm SL), São Paulo, Taubaté, rio Paraíba do Sul, c. 22 ° 58 ’ S, 45 ° 36 ’ W; E. Garbe, 1911. CAS 68912 (1, 161.2 mm SL), “ São Paulo ” (no precise locality); collector unknown, 1902	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F54FFD24EA4FE8BFC07FCF3.taxon	description	Rio de Janeiro, rio Paraíba do Sul basin: MCZ 21114 (1, 135.3 mm SL); MCZ 21115 (4, 130.5 – 149.8 mm SL), “ Mendez ” [rio Paraíba do Sul or tributary at Mendes, 22 ° 32 ' S, 43 ° 44 ' W]; C. F. Hartt & E. Copeland, 1865. MCZ 21110 (7, 154.0 – 178.5 mm SL): “ Muriahé ” [rio Muriaé at Muriaé, 21 ° 8 ' S, 42 ° 23 ' W]; C. F. Hartt & E. Copeland, 1865. MCZ 21117 (5, 148.8 – 231.6 mm SL), “ Campos ” [Rio Muriaé at Campos dos Goytacazes, 3 miles from town, 21 ° 43 ’ S, 41 ° 22 ’ W]; C. F. Hartt & E. Copeland, 1865. CAS 68832 (1, 258.0 mm SL), rio Paraíba do Sul drainage [precise locality uncertain]; Bourget, date not specified [1865 – 1866]. MNRJ 18246 (1, 198.8 mm SL), Itaperuna, Valão de São Domingos (tributary of rio Muriaé, rio Paraíba do Sul basin, 21 ° 16 ' S, 41 ° 46 ' W; D. F. Moraes Jr. et al., 22 Feb 1990. MNRJ 11226 (1, 101.8 mm SL), Itaperuna, córrego do Toiama, trib. Rio Muriaé, road BR- 393 (Bom Jesus do Itabapoana-Santo Antônio de Pádua), c. 21 ° 14 ' 0 '' S, 41 ° 55 ' 30 '' W; D. F. Moraes Jr., G. W. Nunan & L. C. Alvarenga, 15 Jan 1982. MNRJ 4761 (1, 138.3 mm SL), Fazenda da Salgada and São José, between Itaperuna and Miracema; A. L. Carvalho, no date. MZUSP 3043 (1, 236.8 mm SL), Cardoso Moreira, rio Muriaé (tributary of rio Paraíba do Sul), 21 ° 29 ’ S, 41 ° 36 ’ W; O. Pinto, July 1941. MZUSP 103045 (4, 194.5 – 259.0 mm SL): Campos dos Goytacazes, Rio Muriaé, near its mouth, c. 21 ° 42 ’ S, 42 ° 31 ’ W; collector not especified, 2002. MNHN 1909 - 214 (1, 287.7 mm SL): Rio Grande (tributary of rio Paraíba do Sul), c. 21 ° 37 ’ S, 41 ° 49 ’ W; C. Jobert, c. 1878. MZUSP 79088 (2, 141.7 – 148.3 mm SL); MZUSP 79091 (1, 146.5 mm SL): Fish hatchery, Projeto Piabanha, Itaocara (stocked from specimens collected at the middle rio Paraíba do Sul). Rio de Janeiro, coastal river systems: MCZ 21112 (1, 232.2 mm SL), Santa Cruz [rio Grande (Arroio Fundo), in urban Rio de Janeiro, at Fazenda Santa Cruz, 22 ° 53 ' S, 43 ° 43 ' W]; D. Pedro II, leg. D. Bourget, April 1865. MZUSP 88515 (2, 235.0 – 247.0 mm SL), Silva Jardim, Rio São João, near mouth at Lagoa de Juturnaíba, 22 ° 34 ' S, 42 ° 18 ' W; collector not specified, Oct 2005. MZUSP 103029 (1, 291.0 mm SL), Campos dos Goytacazes, rio Imbé, c. 21 ° 47 ’ S, 41 ° 33 ’ W; G. Sousa, 2000. MZUSP 79092 (1, 187.2 mm SL), Fish hatchery at Itaocara (Projeto Piabanha) (stocked from specimens collected at Campos dos Goytacazes, rio Imbé, c. 21 ° 47 ' S, 41 ° 33 ' W); G. Souza, no date. MZUSP 2091 (5, 140.3 – 220.3 mm SL); CAS 11817 (1, 228.0 mm SL); CAS 11816 (1, 213.0 mm SL); Quiçamã, Lagoa Feia, c. 21 ° 57 ’ S, 41 ° 22 ’ W; E. Garbe, Dec 1911. MZUSP 107280 (1, 260.0 mm SL): Campos dos Goytacazes, fish market (probably from Lagoa Feia); O. T. Oyakawa et al., 3 Sept 2004. Rio de Janeiro / Espírito Santo, rio Itabapoana system: MCZ 21116 (9, 144.0 – 191.8 mm SL), Itabapoana [Rio Itabapoana at Itabapoana, 21 ° 17 ’ S, 40 ° 59 ’ W]; C. F. Hartt & E. Copeland, 1865. MZUSP 79087 (3, 193.5 – 231.0 mm SL): rio Itabapoana, c. 21 ° 17 ’ S, 40 ° 59 ’ W; F. Daudt, 14 Aug 2002. Minas Gerais, rio Paraíba do Sul system: ZUEC 6384 (1, 251.0 mm SL): Itamarati de Minas, rio Novo (trib. rio Pomba), 21 ° 28 ’ 36 ’’ S, 42 ° 51 ’ 36 ’’ W; D. M. Rosa, Feb 2011.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F5DFFD64EA4FCA0FD5AFEA7.taxon	diagnosis	Diagnosis. Brycon vermelha can be diagnosed from all remaining cis-andean Brycon species, with the exception of B. stolzmanni, B. coxeyi, B. coquenani, B. insignis, B. howesi, B. dulcis, B. ferox, B. vonoi, B. opalinus, and B. nattereri by possessing a color pattern consisting in a humeral blotch and a caudal peduncle blotch, without body stripes or other obvious color markings on caudal and anal-fins (vs. body stripes and caudal / anal fin color markings present; see Fig. 5). Brycon vermelha can be diagnosed from these species, with the exception of B. howesi, B. insignis, and B. coquenani, by possessing a fifth infraorbital bone considerably wider than high (vs. fifth infraorbital bone about as wide as high; see Fig. 6). Brycon vermelha can be easily distinguished from B. insignis and B. howesi by possessing scarlet-red colored caudal, dorsal, adipose, anal and pelvic-fins in living, clear in preserved specimens (vs. darkened fins, without any red pigmentation in both living and preserved specimens in B. insignis and B. howesi). Brycon vermelha can be distinguished from B. coquenani (from which the color pattern in life is unknown) by possessing a higher head (70.3 – 75.5 % of head length, mean 72.3, vs. 59.8 – 64.9 %, mean 61.4, in B. coquenani), and by possessing pointed penta- to heptacuspidate dentary teeth, with moderately developed lateral cusps (vs. pointed, tri- to tetracuspidate dentary teeth, with lateral cusps poorly developed). See “ Remarks ”, below, for further notes on the diagnosis of the species.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F5DFFD64EA4FCA0FD5AFEA7.taxon	description	Description. Morphometric data are presented in Table 6. Large-sized species, largest examined specimen 395.0 mm SL. Body moderately high in specimens larger than 370 mm SL, moderately slender in specimens between 200 – 230 mm SL. Largest body height immediately ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly to moderately convex from latter point to basis of supraoccipital process, and moderately convex from latter point to dorsal-fin origin, straight along dorsalfin basis, 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 conspicuously acute anteriorly, mouth terminal. Premaxillary and dentary isognathous, i. e., approximatelly of same size. Maxillary long, extending posteriorly to slightly posterior to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Eight (1), 9 (3), 10 (1), 11 (3), 12 (1), or 13 (1) tricuspidate teeth, with poorly developed lateral cusps, in outer series. Four (1), 5 (7), or 6 (2) tricuspidate teeth in second, inner premaxillary row, plus 2 (9) or 3 (1) tri- to tetracuspidate teeth between the first and third rows. Two teeth in third premaxillary row, medial teeth largest, symphyseal teeth smaller, slightly tilted towards each other, both pentacuspidate. Maxillary with distal portion distinctly expanded and rounded in profile. Twenty to 29 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 10 (1), 11 (1), 13 (3), 14 (3), 15 (1), or 16 (1) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, hexa- to heptacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, tri- to unicuspidate. Inner (lingual) series consisting of a small, single unicuspid symphyseal tooth, situated immediately posterior to symphyseal dentary teeth of main series, plus a row of six to eleven small, unicuspidate teeth in the posterior portion of the dentary. Inner symphyseal teeth present in three out of nine specimens (MZUSP 58049, 229.5 mm SL; LBP 9066, 213.0 mm SL) examined for this character. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-seven (1), 51 (2), 52 (4), 53 (2), or 54 (1) scales in lateral line series. Laterosensory tube simple in middle-sized (200 – 230 mm SL) specimens, deflected upwards in the first 5 – 6 scales, downwards in the remaining lateral-line scales. Larger specimens (> 370 mm SL) with branched tubules, mostly bifurcated but some scales with 3 or 4 tubules. Horizontal scale rows between dorsal-fin origin and lateral line 10 (5), 11 (3), or 12 (2). Horizontal scale rows between lateral line and pelvic-fin 4 (3), 5 (4), or 6 (3). Circumpeduncular scales 17 (1), 18 (1), 19 (6), or 20 (1). Dorsal-fin rays ii, 9. Dorsal fin origin at, or slightly ahead of, middle of SL. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 21 (1), 22 (3), 23 (3), or 24 (3). Last unbranched and anterior 3 – 4 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Sheath of scales covering basis of anal-fin rays composed of three scale rows, lower scale row formed by 25 – 30 rectangular scales. Pectoral-fin rays i, 12 (1), 13 (7), or 14 (2). Pelvic-fin rays i, 7. Main caudal-fin rays 10 / 9. Caudal fin forked, lobes slightly pointed. First branchial arch with 8 (1), 10 (2), or 11 (1) lower, 1 at angle, and 8 (2) or 9 (2) upper gill rakers. Coloration in alcohol. Top of head, snout, supraorbital, sixth infraorbital, upper portion of fifth infraorbital, and dorsal portion of body dark- to chestnut-brown. Second, third, fourth and lower portion of fifth infraorbitals silvery hue. Opercle mainly silvery, with some light-brown tinge in specimens which retained relatively less guanine. Dentary, maxillary, gular area and lower portion of body brownish. Lateral portion of body silvery in specimens retaining a considerable amount of guanine, light brown in specimens that lost most guanine (e. g., MZUSP 56048). Humeral blotch present, very little conspicuous, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of second, extending longitudinally to posterior margin of fourth lateral line scales, and vertically less than one scales high. Large, conspicuous, oval-shaped caudal peduncle blotch, extending along 7 – 8 last lateral-line scales. Dorsal, anal, and caudal-fins brownish, with scattered dark pigmentantion on interradial membranes. Pectoral, pelvic and adipose-fins light brown. Coloration in life. Based on pictures of an unpreserved specimen, collected at the rio Mucuri nearby Carlos Chagas, plus color transparencies of the type series. Overall body color silvery. Snout and top of head dark grey. Dorsum, fifth and sixth infraorbital bones plumbeous. Dentary, gular and ventral areas clear. Series of red spots on four to five lateral scales rows situated below lateral line, extending to scales situated at the level of anal-fin origin. Rounded caudal peduncle blotch dark, very conspicuous. Adipose, anal and caudal-fin scarlet-red. Distal margin of dorsal fin scarlet red. Pelvic fins reddish. Pectoral fins hyaline. Sexual dimorphism. Not observed. Hooks were not found in any examined specimen, including the two male paratypes (MZUSP 53304 and USNM 320481), which are in an obvious breeding condition, with fully mature testicles. Common names. “ Vermelha ” (Lima & Castro, 2000: 159).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F5DFFD64EA4FCA0FD5AFEA7.taxon	distribution	Distribution. Known from the middle and upper portions of the rio Mucuri basin, a coastal river system draining northern Minas Gerais and southern Bahia states, Brazil (Fig. 25). The species is apparently absent from the lower portion of this river system (Pompeu & Vieira, 2008). Fishermen from Carlos Chagas report the occurrence of the species at the rio São Mateus, an independent small coastal system, situated immediately south to the rio Mucuri at the Espírito Santo state, but its occurrence at this basin was so far not confirmed (L. F. S. Ingenito & L. F. Duboc, pers. comm.). Ecological notes. All type specimens were collected at a river stretch within a forest remnant. Fishermen of the rio Mucuri report that, in fact, Brycon vermelha favor areas with preserved riparian forest. Stomach contents of the the paratypes included a loricariid catfish, a Rhamdia sp. of about 95 mm SL, and unidentified fruit remains (Lima & Castro, 2000: 161). Both paratypes were males, with fully mature testicles, and were evidently in breeding condition. The species is more common in the middle and lower stretches of the rio Mucuri, where it represents 2 % of the specimens captured in experimental fishing (Pompeu & Vieira, 2008). In contrast, at the area of Santa Clara hydroelectric dam, at the divide between Minas Gerais and Bahia states and at the transition between the middle and lower rio Mucuri, out of 45.000 fish specimens that bypassed the fish ladder during the fish migration season at 2003 / 2004, only two were Brycon vermelha (Pompeu & Vieira, 2008). Brycon vermelha occurs syntopically with the congener B. ferox (see “ Ecological notes ” of this species).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F5DFFD64EA4FCA0FD5AFEA7.taxon	description	Conservation. Brycon vermelha is officially considered threatened with extinction in Brazil (Pompeu & Vieira, 2008). Causes for concern are the limited natural distribution of the species, restricted to a single small hydrographic basin in eastern Brazil, coupled with its preference for river stretches with preserved forest remnants in an area which is now mostly reduced to pastureland. Additionally, a small hydroelectric dam, the Mucuri dam, was build right at the core area of occurrence of the species in the upper rio Mucuri near Carlos Chagas. There is no recent information as to the conservation status of Brycon vermelha after the completion of this dam but with all likelihood the species has become even more imperiled than was before this event.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F5DFFD64EA4FCA0FD5AFEA7.taxon	discussion	Remarks. The scarlet-red fin coloration displayed by Brycon vermelha is a unique feature that promptly diagnoses the species from all remaining Brycon species occurring in the coastal river drainages of eastern Brazil. Among the Brycon species occurring in the coastal river drainages from eastern Brazil possessing an acute head profile, Brycon vermelha is more similar to B. insignis and B. howesi. Brycon vermelha is very similar to B. insignis, and there are apparently no unambiguous features that allow their diagnosis other than the fin color. Brycon vermelha, however, possess more volumous dentaries and premaxillaries than B. insignis, which results in a more rounded snout and mandible profile than in the former. Also, dentary and premaxillaries are about the same size in Brycon vermelha, while in most individuals of B. insignis the premaxilla extends a little beyond the dentary, i. e., the jaws are anisognathous. Body color pattern in Brycon vermelha differ at least from the wild-caught B. insignis examined by lacking the striped and reticulated dark pigmentation patterns present in the latter, as well as for displaying a humeral blotch much less conspicuous than the one displayed by B. insignis. Gill raker counts also seem to differ between both species: Brycon vermelha possess 8 – 11 lower, and 8 – 9 upper gill rakers, vs. 11 – 16 lower, and 11 – 17 upper gill rakers in B. insignis. For a comparison between Brycon vermelha and B. howesi, see item “ Diagnosis ” of the latter species. Though Brycon vermelha was only recently described, a specimen of the species was already collected by Charles Hartt and Edward Copeland during the Thayer Expedition, between 1865 – 1866 (MCZ 162613). This specimen was collected in the rio Mucuri at Santa Clara, a portion of this river where the species is now thought to be naturally scarce (see “ Ecological notes ”, above). Interestingly, Steindachner, who studied in detail material collected by the Thayer Expedition in eastern brazilian rivers, and described some of the Brycon species from eastern Brazil based on it (Steindachner, 1877), did not noticed this particular specimen.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F5DFFD64EA4FCA0FD5AFEA7.taxon	materials_examined	Material examined. Type material: MZUSP 53303 (1, 395.0 mm SL), Brazil, Minas Gerais, Carlos Chagas, rio Mucuri, rio Mucuri, approximately 9 km W of village of Presidente Pena, along a dirt road on Fazenda Gavião, c. 17 ° 37 ’ S, 40 ° 55 ’ W; R. M. C. Castro & S. L. Jewett, 17 – 23 July 1991. Holotype of Brycon vermelha. MZUSP 53304 (1, 372.0 mm SL); USNM 320481 (1, 379.5 mm SL), same data as holotype; paratypes. Non types. Brazil, Minas Gerais: UFMG uncataloged (1, 223.5 mm SL), Carlos Chagas, rio Mucuri; V. Vono, no date. MZUSP 58049 (1, 229.5 mm SL), Carlos Chagas, rio Mucuri; V. Vono, 14 Jan 1997. MZUSP 70217 (2, 215.3 – 233.6 mm SL), Carlos Chagas, rio Mucuri, above Presidente Pena village; F. Vieira, April 2001. LBP 9066 (2, 213.0 – 255.0 mm SL): Minas Gerais, rio Mucuri, c. 17 ° 41 ’ S, 40 ° 46 ’ W; J. A. Senhorini, 6 Feb 2010. Bahia: MCZ 162613 (1, 203.0 mm SL): Santa Clara, rio Mucury [Rio Mucuri at Santa Clara], c. 17 ° 54 ’ S, 40 ° 13 ’ W; C. F. Hartt & E. Copeland, Dec 1865 – April 1866.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F59FFCA4EA4FE53FB7CF9CB.taxon	description	Chalceus opalinus (non Cuvier): Valenciennes, in Cuvier & Valenciennes, 1850: 246 (part; “ Rio Tiquilelonha ”).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F59FFCA4EA4FE53FB7CF9CB.taxon	description	Brycon devillei (non Castelnau): Vieira et al., 2008: 47 – 48 (part; Rio Jequitinhonha basin; ecology, conservation); Azevedo et al., 2011: 807 – 814 (feeding ecology and social behavior, rio Preto, rio Jequitinhonha basin, Minas Gerais). Brycon sp.: Pugedo et al., 2016: 345 (photo), 346 (table) (barcoding, rio Jequitinhonha).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F59FFCA4EA4FE53FB7CF9CB.taxon	diagnosis	Diagnosis. Brycon howesi can be diagnosed from all remaining cis-andean Brycon species, with the exception of B. stolzmanni, B. coxeyi, B. coquenani, B. vermelha, B. insignis, B. dulcis, B. ferox, B. vonoi, B. opalinus, and B. nattereri by possessing a color pattern consisting in a humeral blotch and a caudal peduncle blotch, without body stripes or other obvious color markings on caudal and anal-fins (vs. body stripes and caudal / anal fin color markings present; see Fig. 5). Brycon howesi can be diagnosed from these species, with the exception of B. insignis, B. vermelha, and B. coquenani, by possessing a fifth infraorbital bone considerably wider than high (vs. fifth infraorbital bone about as wide as high; see Fig. 6). Brycon howesi can be diagnosed from B. coquenani by possessing a higher head (65.6 – 76.2 % of head length, mean 70.6, vs. 59.8 – 64.9 %, mean 61.4, in B. coquenani), by possessing pointed tri- to pentacuspidate dentary teeth, with moderately developed lateral cusps (vs. pointed, trito tetracuspidate dentary teeth, with lateral cusps poorly developed), and by mature males displaying hooks only in anal and pelvic-fins (vs. mature males displaying hooks in all rayed fins, including caudal fin). Brycon howesi can be distinguished from B. vermelha by presenting darkened dorsal, caudal, and anal-fins in both living and preserved specimens (vs. dorsal, caudal, and anal-fins reddish in living, clear in preserved B. vermelha specimens), 7 – 9, modally 8 horizontal scale rows between the dorsal-fin origin and the lateral line (vs. 10 – 12, modally 10, in B. vermelha). Brycon howesi can be distinguished from Brycon insignis by possessing 46 – 50, modally 49 lateral line scales (vs. 51 – 65, modally 58 in B. insignis), 7 – 9, modally 8 horizontal scale rows between the dorsal-fin origin and the lateral line (vs. 9 – 11, modally 10 – 11 in B. insignis), 3 – 5, modally 4 horizontal scales rows between the lateral line and pelvic (vs. 4 – 7, modally 5 in B. insignis) and a relatively rounder snout profile in B. howesi (vs. snout more pointed in B. insignis).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F59FFCA4EA4FE53FB7CF9CB.taxon	description	Description. Morphometric data are presented in Table 7. Large-sized species, largest alcohol-preserved examined specimen 284.0 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, 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 pelvicfin 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 considerably acute anteriorly, mouth terminal. Jaws slightly anisognathous, premaxillary projecting slightly relative to dentary, outer row of premaxillary teeth exposed when mouth is closed in some specimens. Maxillary long, extending posteriorly to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Five (1), 6 (1), 7 (3), 8 (10), 9 (9), 10 (8), 11 (4), or 12 (1) tricuspidate teeth in outer series. Four (4), 5 (18), 6 (11), or 7 (3) tri- to tetracuspidate teeth in second, inner premaxillary row, plus 1 (2), 2 (15), 3 (17), or 4 (2) 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, both pentacuspidate. Maxillary with distal portion considerably expanded and rounded in profile. Seventeen to 33 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 8 (1), 10 (3), 11 (5), 12 (8), 13 (3), 15 (2), 16 (1), or 17 (1) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, tetra- to pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, tetra, tri- to unicuspidate. Main dentary present gaps (diastemas) between teeth in small specimens (MZUSP 5139, 6, 31.3 – 47.6 mm SL). Inner (lingual) series consisting of a small, single unicuspid symphyseal tooth, situated immediately posterior to symphyseal dentary teeth of main series, plus row of 23 – 25 small, unicuspidate teeth, originating on lingual crest of dentary replacement trench at level of the fifth to sixth teeth of main series. Inner symphyseal teeth present in all specimens with relatively intact symphyseal dentary area. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-four (1), 45 (4), 46 (6), 47 (7), 48 (10), 49 (7), or 50 (2) scales in lateral line series. Laterosensory tube simple in relatively small (<225 mm SL) specimens, deflected upwards in the first 5 – 6 scales, downwards in the remaining lateral-line scales. Larger specimens (> 225 mm SL) with some scales with 2 – 3 branches, most lateral scales with a single branch. Tubules short, generally not extending beyond scale anterior third. Horizontal scale rows between dorsal-fin origin and lateral line 7 (2), 8 (31), or 9 (4). Horizontal scale rows between lateral line and pelvic-fin 3 (5), 4 (30), or 5 (2). Circumpeduncular scales 14 (3), 15 (16), 16 (15), or 17 (3). Dorsal-fin rays ii, 9. Dorsal fin origin at, or slightly ahead of, middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 12 th (2) or 13 th (1) vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 19 (1), 20 (1), 21 (1), 22 (4), 23 (9), 24 (9), 25 (8), or 26 (1). First anal-fin pterygiophore inserting behind haemal spine of 22 th (1), 23 th (1), or 24 th (1) vertebrae. Last unbranched and anterior 3 – 4 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 20 per fin-ray main branch) middle-sized hooks on last unbranched and posterior main branch of branched rays 17 – 23, associated with dense, gelatinous tissue in six specimens (MZUSP 55963, 221.8 mm SL; MZUSP 101506, 3, 189.2 – 219.0 mm SL; MZUSP 101507, 1, 186.1 mm SL; ZUEC 12869, 245.0 mm SL). A single hook per ray segment. Sheath of scales covering basis of anal-fin rays composed of two scale rows, lower scale row formed by 20 – 25 rectangular scales. Pectoral-fin rays i, 11 (1), 12 (11), 13 (18), 14 (4), or 15 (1). Pelvicfin rays i, 7, two specimens i, 8. A few small hooks on posterior branch of branched pelvic-fin rays in five specimens (MZUSP 55963, 221.8 mm SL; MZUSP 101506, 3, 189.2 – 219.0 mm SL; MZUSP 101507, 1, 186.1 mm SL). Main caudal-fin rays 10 / 9. Caudal fin forked, lobes slightly pointed. Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First branchial arch with 11 (2), 12 (1), 13 (1), 14 (5), 15 (1), or 16 (1) lower, 1 at angle, and 10 (3), 11 (4), or 12 (4) upper gill rakers. Vertebrae 42 (1) or 43 (2). Supraneurals 9 (1) or 10 (2). Coloration in alcohol. Overall color pattern clear in specimens from most localities, but very dark in specimens from the rio Preto (MZUSP 101506, MZUSP 101507, MZUSP 101508). Top of head, snout, supraorbital, sixth infraorbital, and dorsal portion of body brown to dark-brown. Remaining infraorbitals, opercle, dentary, maxillary, lateral and ventral surfaces of body light-brown to brown. Specimens retaining guanine (e. g., MZUSP 55963, ZUEC 12869) with infraorbitals, opercle, and lateral surfaces of body with a silvery hue. Humeral blotch present, little to moderately conspicuous, situated at level of lateral line, extending longitudinally from second to fourth, and vertically to scale row situated immediately above lateral line. Caudal peduncle blotch moderately to highly conspicuous, aproximately rounded in shape, extending along 6 – 7 last lateral line scales. Pectoral, pelvic, dorsal, and anal-fins clear, with a moderate concentration of dark chromatophores at the interradial membranes in the clear-colored specimens; intensely dark-pigmented in the specimens from rio Preto (MZUSP 101506, MZUSP 101507, MZUSP 101508). Caudal fin with outer caudal fin rays with dark pigmentation on outer rays and basis of middle cudal-fin rays continuous with caudal peduncle blotch, forming a roughly V-shaped blotch. Adipose fin light- to dark-brown. Juveniles (MZUSP 5139, 31.3 – 47.6 mm SL) presenting a conspicuous humeral blotch and caudal blotch extending into 3 – 4 innermost caudal-fin rays. Color in life. Based on pictures of freshly collected or living specimens from the rio Preto, Rio Preto State Park, taken by P. Azevedo, a specimen collected at the rio Vacaria, a tributary of the rio Jequitinhonha, by F. R. Andrade Neto, and several specimens from the rio Itacambiruçu, collected by T. C. Pessali. Overall color pattern clear-colored to dark-brown; clear-colored specimens with a silvery hue on opercle and infraorbital bones and lateral surfaces of body. Caudal peduncle blotch and dark pigmentantion on outer caudal-fin rays very conspicuous. Adipose-fin cream-colored to orangish. Specimens from the clear to turbid water rio Vacaria and rio Itacambiruçu possess an overall color pattern much more clear when compared with specimens from the black-water rio Preto. Sexual dimorphism. Five specimens (MZUSP 55963, 221.8 mm SL; MZUSP 101506, 3, 189.2 – 219.0 mm SL; MZUSP 101507, 1, 186.1 mm SL) were recorded as possessing anal- and pelvic-fin hooks, a single specimen (ZUEC 12869, 245.0 mm SL) presented only anal-fin hooks. One specimen (MZUSP 101507, 1, 186.1 mm SL) proved to be a male after dissection, with relatively well-developed testes. There is an indication in the literature that females may grown larger than males (see “ Ecological notes ”, below).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F59FFCA4EA4FE53FB7CF9CB.taxon	etymology	Etymology. Named in honour of Gordon J. Howes (1938 – 2013), in recognition to his important contribution to the knowledge of the taxonomy of the genus Brycon as well as for ichthyology as a whole. Common names. “ Piabanha ” (Godinho et al., 1998: 415; V. Vono, P. Azevedo, T. C. Pessali, pers. comm.). Valenciennes (in Cuvier & Valenciennes, 1850: 246) mentions the common name “ pirabanha ” for the species, a name apparently heard by A. de Saint-Hillaire, the collector of the specimen studied by him.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F59FFCA4EA4FE53FB7CF9CB.taxon	distribution	Distribution. Endemic from the rio Jequitinhonha basin, an independent coastal river system at the state of Minas Gerais, Brazil (Fig. 25). Ecological notes. Feeding and social behavior of the species was quantitavely studied by Azevedo et al. (2011; as B. devillei) at a dark water tributary of the rio Araçuaí (a tributary of the rio Jequitinhonha). Brycon howesi adopted mainly surface-picking as a foraging tactic, though “ digging ” and “ active-hunting ” were also observed (Azevedo et al., 2011). Surface-picking was more common during the rainy season, whereas “ digging ” was more frequent during the dry season (Azevedo et al., 2011). An elaborate cooperative hunting of small characids, and a feeding association of the kind “ nuclear-follower behavior ” with Leporinus garmani, a bottom feeder, were also documented for the species (Azevedo et al., 2011). Brycon howesi was reported by Godinho et al. (1998; as B. insignis) as being mainly insectivore. Largest recorded standard lengths reported for the species by these authors were 254.0 mm SL for males and 342.0 mm SL for females. The species occurs both in clear- (e. g., rio Itacambiruçu) and dark-water (e. g., rio Preto) tributaries of the rio Jequitinhonha basin.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F59FFCA4EA4FE53FB7CF9CB.taxon	description	Conservation. The rio Jequitinhonha basin has a long history of anthropogenic disturbances that includes intensive mining, deforestation, and siltation. Probably as a result of these impacts, Brycon howesi was reported by fishermen to be becoming increasingly scarcer in the middle rio Jequitinhonha basin (Godinho et al., 1998). A large hydroelectric dam, the Irapé dam, was build in 2006 a little above the area where most of the records of Brycon howesi come from, in the middle rio Jequitinhonha basin, and the extent of its presumable deleterious effects on the populations of the species are still unknown. Fortunately, an apparently healthy population of the species occurs at the Rio Preto State Park, at the headwaters of the rio Araçuaí, the main tributary of the rio Jequitinhonha (Azevedo et al., 2011).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F59FFCA4EA4FE53FB7CF9CB.taxon	discussion	Remarks. There are two references to specimens of Brycon howesi in the taxonomic literature, both as misidentifications. The first report for the species was by Valenciennes (in Cuvier & Valenciennes, 1850: 246), which identified it as Chalceus opalinus (now Brycon opalinus; see under this species). This record was based on a single, stuffed specimen (MNHN A. 8612), collected by the botanist Auguste de Saint-Hilaire at the rio “ Tiquilelonha ” (an evident misspelling of Jequitinhonha), a region surveyed by him between April – July 1817 (Papavero, 1971). This specimen certainly does not belong to the type-series of Chalceus opalinus, which is composed only of the holotype, MNHN A. 8613 (see item “ Remarks ” of Brycon opalinus, below). Although Géry & Mahnert (1992: 813) considered the specimen MNHN A. 8612 to be probably conspecific with the holotype of Chalceus opalinus, our examination of this same specimen leaves no doubt that it in fact represents Brycon howesi. Steindachner (1877: 591) reported “ Rio Jequitinhonha ” as being one of the syntypical localities of Brycon insignis, the remaining localities (“ Mendez ” and “ Campos ”) being situated at the rio Paraíba do Sul basin. A search in the NMW fish collection revealed a single specimen, NMW 62927 (1, 71.6 mm SL) collected at the “ Rio Jequitinhonha ”, which undoubtedly represents the material reported by Steindachner (1877). The designation of the specimen NMW 62925 (144.8 mm SL) as the lectotype of Brycon insignis resulted in restricting this name for the species occurring in the rio Paraíba do Sul drainage (see “ Remarks ” of Brycon insignis). Thus, the specimen NMW 62927, though a paralectotype of Brycon insignis, is not conspecific with the lectotype and the remaining paralectotypes, representing instead the species which is herein named as Brycon howesi. In fact, Brycon howesi is more similar to B. insignis than to any other Brycon species. The V-shaped caudal-fin blotch formed by pigmentation concentrated on the outer caudal-fin rays is a feature uniquely shared between these two species. As noticed in the item “ Diagnosis ”, above, Brycon howesi is clearly distinct from B. insignis by possessing significantly lower scale counts and a distinctly rounder snout profile.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F59FFCA4EA4FE53FB7CF9CB.taxon	materials_examined	Material examined. Holotype: MZUSP 103075 (230.6 mm SL): Brazil, Minas Gerais, Araçuaí, rio Jequitinhonha, confluence with rio Araçuaí, below village of Itira, 16 ° 45 ’ S, 42 ° 0 ’ W; J. C. Garavello, A. S. Soares, A. I. Alves & J. C. Soares, 19 Feb 1989. Paratypes: Brazil, Minas Gerais, rio Jequitinhonha basin: MZUSP 53803 (1, 234.8 mm SL): same data as holotype. MZUSP 53802 (1, 223.1 mm SL): Leme do Prado, rio Araçuaí at Santa Rita village, 17 ° 6 ’ S, 42 ° 39 ’ W; G. B. Santos, A. S. Soares & A. I. S. Alves, 18 Oct 1987. MZUSP 55963 (6, 165.8 – 248.6 mm SL); ZUEC 6365 (1, 164.9 mm SL): Coronel Murta, rio Jequitinhonha, 16 ° 37 ’ 32 ’’ S, 42 ° 16 ’ 6 ’’ W; F. Andrade, 6 Aug 2010. MZUSP 5139 (6, 1 cs, 31.3 – 47.6 mm SL), Itaobim, rio Jequitinhonha, 16 ° 34 ’ S, 41 ° 29 ’ W; Exc. Departamento de Zoologia, 25 June 1966. ZUEC 6939 (3, 134.0 – 242.0 mm SL): Grão Mogol, rio Itacambiruçu, 16 ° 34 ’ 52 ’’ S, 42 ° 52 ’ 25 ’’ W; T. C. Pessali & T. F. Teixeira, May 2011. ZUEC 6940 (1, 143.7 mm SL): Grão Mogol, rio Itacambiruçu, 16 ° 35 ’ 12 ’’ S, 42 ° 51 ’ 15 ’’ W; T. C. Pessali & T. F. Teixeira, May 2011. ZUEC 7003 (1, 282.0 mm SL); MCP 48399 (1, 268.0 mm SL); ANSP 200242 (1, 232.0 mm SL): Grão Mogol, rio Itacambiruçú, 16 ° 34 ’ 52 ’’ S, 42 ° 52 ’ 25 ’’ W; T. C. Pessali & A. A. Rodrigues, Feb 2011. ZUEC 7000 (1, 208.0 mm SL): Grão Mogol, rio Itacambiruçú, 16 ° 35 ’ 50 ’’ S, 42 ° 50 ’ 22 ’’ W; T. C. Pessali & A. A. Rodrigues, Feb 2011. ZUEC 12869, 1, 245.0 mm SL, Grão Mogol, Itacambiruçu, rio Itacambiruçu, above Cachoeira Marias das Neves, 16 ° 34 ' 53 '' S, 42 ° 52 ' 26 '' W; T. C. Pessali & A. A. Rodrigues, 25 Dec 2011. ZUEC 12870, 1, 213.0 mm SL, Grão Mogol, Itacambiruçu, rio Itacambiruçu, above Cachoeira Marias das Neves, 16 ° 35 ' 58 '' S, 42 ° 55 ' 37 '' W; T. C. Pessali & A. A. Rodrigues, 28 Dec 2011. MZUSP 101506 (4, 167.1 – 219.0 mm SL): São Gonçalo do Rio Preto, rio Preto (tributary of rio Araçuaí), Rio Preto State Park, 18 ° 6 ’ 47 ’’ S, 43 ° 20 ’ 42 ’’ W; P. G. Azevedo, 8 Oct 2006. MZUSP 101507 (3, 186.1 – 243.0 mm SL): same locality and collector, 5 April 2007. MZUSP 103685 (2 skel., 168.0 – 207.0 mm SL): same locality and collector, July 2009. MZUSP 101508 (3, 244.0 – 284.0 mm SL): same locality; F. Vieira & G. B. Santos, 10 Jan 2006. MZUEL 3931 (1, 130.0 mm SL): same locality; E. T. Grando, 2 Feb 2004.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F59FFCA4EA4FE53FB7CF9CB.taxon	description	Non types. Minas Gerais: LISDEBE uncat. (2, 190.0 – 243.0 mm SL): Coronel Murta, rio Jequitinhonha, fazenda Jatobá, c. 16 ° 37 ’ S, 42 ° 10 ’ W; V. Vono, April 1989. ZUEC 6969 (1, 101.7 mm SL): Grão Mogol, rio Itacambiruçú, 16 ° 35 ’ 12 ’’ S, 42 ° 51 ’ 15 ’’ W; T. C. Pessali & A. A. Rodrigues, Feb 2011. ZUEC 6970 (2, 109.1 – 138.4 mm SL): Grão Mogol, rio Itacambiruçú, 16 ° 35 ’ 12 ’’ S, 42 ° 51 ’ 15 ’’ W; T. C. Pessali & A. A. Rodrigues, Feb 2011. ZUEC 6974 (3, 101.1 – 171.5 mm SL): Grão Mogol, rio Itacambiruçú, 16 ° 34 ’ 52 ’’ S, 42 ° 52 ’ 25 ’’ W; T. C. Pessali & A. A. Rodrigues, Feb 2011. ZUEC 6999 (1, 173.3 mm SL): Grão Mogol, rio Itacambiruçú, 16 ° 34 ’ 24 ’’ S, 42 ° 49 ’ 46 ’’ W; T. C. Pessali & A. A. Rodrigues, Feb 2011. ZUEC 6367 (1, 94.3 mm SL): Grão Mogol, rio Itacambiruçu, 16 ° 35 ’ 56 ’’ S, 42 ° 50 ’ 8 ’’ W; F. Andrade, 22 Nov 2010. Imprecise localities: NMW 62927 (1, 71.6 mm SL): “ Rio Jequitinhonha ” (no specific locality); Wertheimer, no date. Paralectotype of Brycon insignis Steindachner. MNHN A. 8612 (1, 295.8 mm SL): “ Rio Tiquilelonha. A. Saint- Hilaire, 1822 ”.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F45FFCC4EA4F9B8FEB4FE37.taxon	description	Summary information on meristics. Lateral line scales 53. Horizontal scale rows between dorsal-fin origin and lateral line 9. Horizontal scale rows between lateral line and pelvic-fin 4. Circumpeduncular scales 18. Dorsal-fin rays ii, 9. Pectoral-fin rays i, 13. Pelvic-fin rays i, 7. Anal-fin rays iii, 23. Outer premaxillary teeth row with 10 teeth. Four teeth in second, inner premaxillary row, plus 2 teeth between the first and third rows. Fifteen dentary teeth at the main series. Twenty-five maxillary teeth. First branchial arch with 13 lower, 1 at angle, and 12 upper gill rakers. Morphometric data of the holotype presented in Table 8.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F45FFCC4EA4F9B8FEB4FE37.taxon	discussion	Remarks. Brycon devillei is only known from its holotype (MNHN 4517), which has as its stated type-locality “ Bahia ”. Castelnau, who collected and described the species, spent a year as the French consul in Salvador (the capital city of Bahia) in 1849 (Papavero, 1971). “ Bahia ” was often used in the XIX zoological and botanical literature to refer to Salvador and surrounding areas. No Brycon specimens were examined in the present study from the northern coastal river systems adjacent to Salvador, with the exception of a single B. vonoi specimen apparently collected immediately southward from Salvador at the rio Una (see item “ Remarks ” of B. vonoi, below). The holotype of Brycon devillei is distinct from B. vonoi by possessing a distinctly pointed head profile (vs. head profile relatively blunt), a fifth infraorbital bone considerably wider than high (vs. fifth infraorbital bone about as wide as high; see Fig. 6), and a higher number of horizontal scale rows between dorsal-fin origin and lateral line (9, vs. 7 – 8). Overall, the holotype of Brycon devillei is very similar and virtually undistinguishable from B. insignis, a species occurring considerably southward, in the rio Paraíba do Sul and coastal river systems from Rio de Janeiro, eastern Brazil (see item “ Distribution ” of B. insignis, above). Castelnau stayed in Rio de Janeiro during June to October of 1843 (Papavero, 1971) and collected some fishes in the area (e. g., the holotype of Loricariichthys castaneus, a species restricted to coastal river systems from rio Doce basin southward to rio Ribeira de Iguape basin in eastern Brazil; Reis & Pereira, 2000). Kullander & Lucena (2006: 134) remarked that the type-locality of Crenicichla lacustris, said to have been collected at “ Bahia ” by Castelnau (1855), was probably an error, since the species was never collected so far northward into the northern coastal rivers of Bahia, the most setentrional known locality being the rio Buranhém in southern coastal Bahia. They proposed that the holotype of Crenicichla lacustris was more likely collected in the Rio de Janeiro area, where the species is in fact quite common (Kullander & Lucena, 2006: 134). It seems also possible that the holotype of Brycon devillei was collected in the Rio de Janeiro area, instead of Bahia. In that case, Brycon devillei would have to be considered a senior synonym of B. insignis. However, given our current lack of knowledge on the Brycon populations occurring in the coastal river systems north from the rio Pardo, it seems more cautious to await for additional information on the Brycon species occurring in the area, before proposing such a change. To complicate further the issue, at least one specied described by Castelnau (1855) bearing the locality “ Bahia ”, was demonstrated to be based on specimens very likely collected in rivers draining the southern portion of Bahia state (Silva & Malabarba, 2016). Until the matter can be satisfactorily settled, we prefer to consider Brycon devillei as a species inquirenda.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F45FFCC4EA4F9B8FEB4FE37.taxon	materials_examined	Material examined. MNHN 4517 (1, 145.5 mm SL): “ Bahia ”. F. de Castelnau, no date. Holotype of Chalceus devillei Castelnau.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F43FFC14EA4FDE3FD3CF95B.taxon	description	Brycon sp.: Travassos & Freitas, 1948: 628 (Espírito Santo, Lagoa de Juparanã, Brazil: parasites).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F43FFC14EA4FDE3FD3CF95B.taxon	diagnosis	Diagnosis. Brycon dulcis can be diagnosed from all remaining cis-andean Brycon species, with the exception of B. stolzmanni, B. coxeyi, B. coquenani, B. vermelha, B. insignis, B. howesi, B. ferox, B. vonoi, B. opalinus, and B. nattereri by possessing a color pattern consisting in a humeral blotch and a caudal peduncle blotch, without body stripes or other obvious color markings on caudal and anal-fins (vs. body stripes and caudal / anal fin color markings present; see Fig. 5). Brycon dulcis can be distinguished from B. stolzmanni, B. coxeyi, B. vonoi, B. opalinus, and B. nattereri by possessing a distinctly acute head profile (vs. a roughly rounded to slightly acute head profile). Brycon dulcis can be additionally distinguished from B. stolzmanni and B. coxeyi by the absence of a patch of dark pigmentation on the opercle (vs. dark patch of pigmentation present on opercle). Brycon dulcis can be distinguished from B. insignis, B. howesi, B. coquenani, and B. vermelha by possessing a fifth infraorbital bone about as wide as high (vs. fifth infraorbital bone wider than high; see Fig. 6). Brycon dulcis can be distinguished from B. ferox by presenting an approximately isognathous mouth, with premaxillary and dentary mostly overlapping, leaving only part of outer premaxillary series exposed in ventral view in some specimens (vs. mouth distinctly anisognathous, premaxillary pointed, extending beyond dentary, leaving outer series, and often also the second, series of premaxillary teeth exposed in ventral view). Brycon dulcis can be additionally diagnosed from other Brycon species occurring in the rio Doce basin, B. opalinus, by possessing a higher number of anal-fin rays (22 – 27, modally 25, vs. 17 – 23, modally 20). See “ Remarks ”, for additional notes on the recognition of the species.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F43FFC14EA4FDE3FD3CF95B.taxon	description	Description. Morphometric data are presented in Table 9. Large-sized species, largest examined specimen 372.0 mm SL. Body slender to moderately slender. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly concave to straight 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 moderately acute anteriorly, considerably acute in specimens <140 mm SL, mouth terminal. Jaws isognathous to slightly anisognathous, premaxillary projecting slightly relative to dentary in some specimens, leaving outer row of premaxillary teeth exposed when mouth is closed. Maxillary long, extending posteriorly to slightly posterior to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Six (3), 7 (5), 8 (8), 9 (8), 10 (11), 11 (7), or 12 (3) tricuspidate teeth in outer series. Three (3), 4 (21), 5 (13), or 6 (4) tetra- to pentacuspidate teeth in second, inner premaxillary row, plus 1 (1), 2 (17), 3 (18), or 4 (2) 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, both hexacuspidate. Maxillary with distal portion distinctly expanded and rounded in profile. Eighteen to 29 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 9 (1), 10 (5), 11 (2), 12 (7), 13 (7), 14 (5), 15 (5), 16 (5), or 17 (2) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, penta-, tetra-, tri- 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 37 – 44 small, unicuspidate teeth, originating on lingual crest of dentary replacement trench slightly after the symphysis. Inner symphyseal teeth present in all specimens with relatively intact symphyseal dentaries area, except for MZUSP 42627 (2, 127.0 – 138.7 mm SL), where symphyseal teeth are missing. Central cusp distinctly larger and pointed, and teeth presenting diastemas in specimens <140 mm SL. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-eight (3), 49 (1), 50 (1), 51 (9), 52 (6), 53 (8), 54 (6), 55 (5), 56 (1), 57 (3), 58 (1), 60 (2), or 64 (1) scales in lateral line series. Laterosensory tube simple in small (<180 mm SL) specimens, deflected upwards in the first 5 – 6 scales, downwards in the remaining lateral-line scales. Larger specimens (> 180 mm SL) with branched tubules, mostly bifurcated in middle-sized specimens but some scales with 3 – 5 tubules. Horizontal scale rows between dorsal-fin origin and lateral line 8 (3), 9 (27), 10 (14), or 11 (3). Horizontal scale rows between lateral line and pelvic-fin 4 (12), 5 (25), or 6 (10). Circumpeduncular scales 16 (2), 17 (7), 18 (17), 19 (11), 20 (5), 22 (1), or 23 (2). Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead of middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 13 th (1), 14 th (1), or 15 th (1) vertebrae. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 22 (4), 23 (7), 24 (11), 25 (19), 26 (4), or 27 (2). First anal-fin pterygiophore inserting behind haemal spine of 23 th (1) or 24 th (2) vertebrae. Last unbranched and anterior 3 – 4 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 20 – 30 per fin-ray main branch) middle-sized hooks on last unbranched and posterior main branch of branched rays 13 – 22, associated with dense, gelatinous tissue in 10 specimens (MZUSP 17070, 265.1 – 270.1 mm SL; MZUSP 36663, 225.7 – 237.9 mm SL; MZUSP 58911, 267.0 mm SL; MZUSP 106770, 285.0 – 323.0 mm SL; MCZ 20031, 236.5 mm SL; NMW 62947, 337.0 – 372.0 mm SL). Some fin rays with hooks also arranged along anterior ray margin. A single hook per ray segment, except when hooks also present on anterior ray margin. Sheath of scales covering basis of anal-fin rays composed of two scale rows, lower scale row formed by 20 – 22 rectangular scales. Pectoral-fin rays i, 11 (2), 12 (8), 13 (24), 14 (9), or 15 (2). Pelvic-fin rays i, 7. Main caudal-fin rays 10 / 9. Caudal fin forked, lobes pointed. Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First branchial arch with 10 (1), 11 (2), 12 (2), 13 (5), 14 (6), or 15 (3) lower, 1 at angle, and 9 (1), 10 (1), 11 (9), 12 (5), 13 (3), or 14 (1) upper gill rakers. Vertebrae 44 (3). Supraneurals 9 (1) or 10 (2). Coloration in alcohol. Overall body coloration clear to dark. Top of head, snout, supraorbital, sixth infraorbital, and dorsal portion of body light to dark-brown. Second, third, and fourth infraorbitals, and opercle light brown, with a silvery hue in specimens retaining guanine (e. g., MZUSP 58911, NMW 62947, ZUEC 6363). Dentary, maxillary, gular area and lower portion of body cream to light brown. Lateral portion of body lightbrown, silvery in specimens retaining a considerable amount of guanine (e. g., MZUSP 58911, NMW 62947, ZUEC 6363). Humeral blotch present, little 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 fourth to fifth lateral line scales, and vertically less than one scale high. Large, moderately conspicuous, oval-shaped caudal peduncle blotch, extending along 8 – 10 last lateral-line scales. All rayed fins with a considerable amount of dark pigmentation at the interradial membranes, imparting an overall dark coloration to these fins. Caudal-fin with a poorly discernible, roughly V-shaped blotch, formed by dark pigmentantion situated on outer caudal-fin rays. Adipose fin light-grey to light-brown. Specimens collected in lakes surrounded by forest, as those from the lakes of the middle and lower Rio Doce (e. g., MZUSP 36663, MZUSP 58911, MZUSP 17070), with a dark overall coloration, while specimens collected in tributaries of the rio Doce in more turbid water conditions (e. g., ZUEC 6776, ZUEC 6363) with a considerably clearer overall coloration. Color in life. Description based on a photo of a freshly collected specimen, collected at Rio Doce State Park by P. S. Pompeu, two specimens (ZUEC 6363, and ZUEC 6776) collected by T. C. Pessali from rio Corrente Grande, and specimens collected at a tributary of rio Santo Antônio and at the upper rio Doce, Rio Doce county (photos supplied by J. Dergam). Top of head, snout, and dorsum light-grey to dark. Dentary and maxillary clear. Opercle, infraorbital bones, and lateral portion of body silvery, with some dark pigmentation in the specimen from the Rio Doce State Park. Humeral blotch inconspicuous; caudal peduncle blotch moderately conspicuous. Series of pinkish spots situated along the four horizontal scale rows situated below scale row immediately below lateral line and anterior to insertion of pelvic fins in the specimens from Rio Doce State Park, upper Rio Doce and rio Santo Antônio. Pinkish pigmentation concentrated on the basal portion of the scales. All fins with some amount of dark pigmentation. Caudal fin with outer fin-rays dark, forming a roughly V-shaped blotch. Sexual dimorphism. As discussed above, anal-fin hooks are present in 10 specimens (MZUSP 17070, 265.1 – 270.1 mm SL; MZUSP 36663, 225.7 – 237.9 mm SL; MZUSP 58911, 267.0 mm SL; MZUSP 106770, 285.0 – 323.0 mm SL; MCZ 20031, 236.5 mm SL; NMW 62947, 337.0 – 372.0 mm SL). Two of them (MZUSP 17070, 270.1 mm SL and MZUSP 36663, 237.9 mm SL) were dissected and are males, with poorly developed testicles. On other hand, a single dissected specimen that does not possess anal-fin hooks (MZUSP 36663, 283.1 mm SL) proved to be a female, with moderately well-developed ovaries.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F43FFC14EA4FDE3FD3CF95B.taxon	etymology	Etymology. Dulcis, after the Latin word for sweet (“ doce ” in Portuguese), in allusion to the river system from which the species is apparently endemic. Common names. “ Piabanha ”.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F43FFC14EA4FDE3FD3CF95B.taxon	distribution	Distribution. Only known from the middle and lower rio Doce basins, states of Minas Gerais and Espírito Santo, eastern Brazil (Fig. 32). There is a putative historical record for the species for the rio Santa Maria da Vitória, a small coastal drainage situated southward from the rio Doce basin (MZUSP 1748, CAS 11171, CAS 11173). However, there is some uncertainty about the exact locality of these lots (though actually the lots from CAS originated from the lot from MZUSP, and as such refer to a single collection event) since it was recorded in the original MZUSP book catalog as being collected in “ Porto Cachoeiro, rio Doce ”. “ Porto Cachoeiro ” is the old name of Santa Leopoldina, a small town situated at the banks of the rio Santa Maria da Vitória, rather than the rio Doce. The collector, Ernst Garbe, collected also at the rio Doce during the same field expedition (Pinto, 1945: 17), so the mixing of contradictory information in the label of this lot does not lend much confidence as to the exact provenance of these specimens. Consequerntly, this locality was not mapped. Ecological notes. Brycon dulcis is recorded as occurring in lake systems associated with the rio Doce, as the Lagoa Juparanã and some other lakes on the lower rio Doce basin and the Lago Dom Helvécio and Lagoa Carioca on the middle rio Doce. Lakes of the middle rio Doce basin, where most the recent records of the species come from, are ancient tributaries of the rio Doce which were dammed either as a consequence of paleoclimatic flutuations or neotectonic activity during the Quaternary (Suguio & Kohler, 1992). It is unclear whether populations of Brycon dulcis occurring in these lakes are in fact isolated from populations in the rio Doce mainstream or not. Stomach contents of two specimens were examined. One specimen (MZUSP 1531, 146.6 mm SL) ingested nine Astyanax sp. (Characidae), one Geophagus brasiliensis (Cichlidae), one Poecilia vivipara (Poeciliidae) and a Lepidoptera larvae. The stomach of the second specimen (MZUSP 36643, 277.8 mm SL) contained unidentified vegetal matter and crushed insects.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F43FFC14EA4FDE3FD3CF95B.taxon	description	Conservation. The rio Doce basin is highly disturbed environmentally, water pollution, siltation and deforestation being the main anthropogenic disturbances in this river basin. Brycon dulcis was apparently extirparted from the lower portion of the basin, in the Espírito Santo state, where it was relatively common until at least the decade of 1960. Recent records of the species come mostly from the natural lakes system at the middle rio Doce in the Rio Doce State Park (Vieira et al., 2008; Gomes et al., 2007; as Brycon devillei). Introduction of two predatory fishes, a species of Cichla and the red-bellied piranha, Pygocentrus nattereri, have however impacted native fish populations in these isolated lakes (Godinho & Formagio, 1992; Godinho et al., 1994). The species still persists at the rio Corrente Grande, a tributary of the rio Doce, in the river stretch between Naque and Periquito, where fishermen report that about three specimens are fished per year (T. C. Pessali, pers. comm.), and was also recently recorded at the upper Rio Doce at Rio Doce county and at the mouth of rio Tanque, a tributary of rio Santo Antônio, near Ferros (J. Dergam, pers. comm., March 2012). The species is considered officially threatened in Brazil (Vieira et al., 2008; as Brycon devillei). The recent catastrophic burst of a tailings dam at the upper rio Doce basin in November 2015, releasing millions of tons of iron waste into the rio Doce, very likely extirpated Brycon dulcis from the rio Doce main channel, though the species might still persist in some tributaries and in the lakes of the middle rio Doce.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F43FFC14EA4FDE3FD3CF95B.taxon	discussion	Remarks. Brycon dulcis was previously misidentified in the literature as B. devillei (e. g., Amaral-Campos, 1950; Sunaga & Verani, 1991; Godinho et al., 1994; Vieira et al., 2008; Travenzoli et al., 2015; see item “ Remarks ” of Brycon devillei and B. ferox). As noticed under Brycon devillei, this latter name is actually a species inquirenda, and more similar to B. insignis. Brycon dulcis is very similar morphologically to B. ferox and B. vonoi. Although diagnostic features among these three species are slight and refer only to overall color pattern and head / snout shape, they are consistent and no gradation among these three morphological types was observed. See additional notes on the item “ Remarks ” of Brycon vonoi. Recently, Travenzoli et al. (2015) argued in favor for the usage of the name Brycon devillei for the Brycon species from the rio Doce herein described as Brycon dulcis. They argued that the species also occurs at the rio Mucuri, which has its mouth at the southern portion of Bahia state, and consequently that the name B. devillei could be applied to the species. We have not examined the specimen from the rio Mucuri that Travenzoli et al. (2015) considered to be conspecific with specimens from the rio Doce and consequently cannot confirm their claim. All Brycon specimens from the rio Mucuri examined in the present study were either B. ferox or B. vermelha (see under these species). At any rate, the name Brycon devillei cannot be applied to the Brycon species herein named B. dulcis because its holotype presents a sixth infraorbital bone wider than high (vs. a sixth infraorbital bone as high as wide in B. dulcis). See also the item “ Remarks ” of Brycon devillei.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F43FFC14EA4FDE3FD3CF95B.taxon	materials_examined	Material examined. Holotype: MZUSP 58911 (1, 267.0 mm SL): Brazil, Minas Gerais, Marliéria, Lago Dom Helvécio, Rio Doce State Park, rio Doce system, 19 ° 46 ’ S, 42 ° 36 ’ W; A. L. Godinho & P. S. Pompeu, Sept 1993. Paratypes: Brazil, Minas Gerais, rio Doce basin: MZUSP 28983 (2, 1 cs, 113.8 – 223.9 mm SL): Marliéria, Lagoa Carioca, Rio Doce State Park, 19 ° 45 ’ S, 42 ° 37 ’ W; J. R. Verani, 27 June – 14 July 1983. MZUSP 36663 (5, 211.3 – 287.9 mm SL): Marliéria, Lago Dom Helvécio, Rio Doce State Park, 19 ° 46 ’ S, 42 ° 36 ’ W; J. R. Verani, 27 Nov – 13 Dec 1985. MZUSP 28968 (1, 111.9 mm SL): Marliéria, Lago Dom Helvécio, Rio Doce State Park, 19 ° 46 ’ S, 42 ° 36 ’ W; J. R. Verani, 25 – 28 Jun 1983. MZUSP 36643 (3, 1 skel., 147.1 – 277.8 mm SL): Marliéria, Lagoa Carioca, Rio Doce State Park, 19 ° 45 ’ S, 42 ° 37 ’ W; J. R. Verani, 23 – 25 Nov 1985. LISDEBE uncat. (7, 245.0 – 260.0 mm SL): Marliéria, Lago Dom Helvécio, Rio Doce State Park, 19 ° 46 ’ S, 42 ° 36 ’ W; J. G. da Silva, March 1977. ZUEC 6363 (1, 263.0 mm SL): Governador Valadares, rio Corrente Grande (trib. rio Doce), 18 ° 59 ’ 38 ’’ S, 42 ° 13 ’ 33 ’’ W; T. C. Pessali, May 2010. ZUEC 6776 (1, 156.0 mm SL): Periquito, rio Corrente Grande (marginal lagoon), 18 ° 58 ’ 20 ’’ S, 42 ° 17 ’ 13 ’’ W; T. C. Pessali, July 2011. Espírito Santo, rio Doce basin: MNRJ 11248 (4, 87.7 – 152.6 mm SL): Linhares, lagoa de Juparanã, road Linhares / São Mateus, 19 ° 16 ’ S, 40 ° 8 ’ W; L. Travassos & H. Travassos, 8 March 1948. MNRJ 11250 (2, 112.8 – 126.9 mm SL): Linhares, lagoa de Juparanã, 19 ° 16 ’ S, 40 ° 8 ’ W; L. Travassos & H. Travassos, 29 Feb 1948. MZUSP 17070 (3, 265.1 – 283.1 mm SL): Linhares, Lagoa Juparanã, 19 ° 16 ’ S, 40 ° 8 ’ W; H. A. Britski & Izáurio Dias, 3 – 9 Feb 1965. MZUSP 3308 (1, 183.3 mm SL): Linhares, rio São José, tributary of Lagoa Juparanã, 19 ° 10 ' S 40 ° 12 ' W; O. Pinto, Sept 1942. Non types. Minas Gerais: MCZ 20031 (3, 194.4 – 236.5 mm SL): Minas Gerais (?), Rio Doce (no specific locality); C. F. Hartt & E. Copeland, Aug – Sept 1865. MZUSP 42627 (2, 127.0 – 138.7 mm SL): Viçosa, c. 20 ° 43 ’ S, 42 ° 54 ’ W; J. Dergam, no date. MZUSP 106770 (2, 285.0 – 323.0 mm SL); MZUSP 106771 (2, 1 skel., 308.0 – 326.0 mm SL): Marliéria, Lagoa Carioca, Rio Doce State Park, 19 ° 45 ’ 26 ’’ S, 42 ° 37 ’ 6 ’’ W; E. N. Fragoso et al., March 2007 – Sept 2009. Espírito Santo: MNRJ 11249 (6, 87.0 – 104.9 mm SL): Linhares, Lagoa Juparanã ,, 19 ° 16 ’ S, 40 ° 8 ’ W; L. Travassos, H. Travassos & J. T. de Freitas, 8 March 1948. MNRJ 11251 (1, 110.7 mm SL): Linhares, Lagoa de Freitas; L. Travassos & J. T. de Freitas, Oct 1963. MZUSP 1531 (5, 1 cs, 88.7 – 146.6 mm SL): Linhares, rio Doce, c. 19 ° 24 ’ S, 40 ° 3 ’ W; E. Garbe, 1906. MCZ 60931 (2, 109.9 – 167.4 mm SL); MCZ 21102 (2, 148.6 – 184.4 mm SL); MCZ 21101 (1, 357.0 mm SL): Rio Doce, between Linhares and Aimorés, 19 ° 37 ’ S, 39 ° 49 ’ W; C. F. Hartt & E. Copeland, 1865. MZUSP 1748 (7, 1 cs, 74.6 – 138.1 mm SL); CAS 11171 (1, 132.4 mm SL); CAS 11753 (1, 105.1 mm SL): Santa Leopoldina, rio Santa Maria da Vitória, 20 ° 6 ’ S, 40 ° 31 ’ W; E. Garbe, 1906.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F43FFC14EA4FDE3FD3CF95B.taxon	description	Rio Doce, locality not specified: NMW 69928 (3, 132.9 – 200.0 mm SL); NMW 62940 (4, 191.5 – 235.0 mm SL); NMW 62947 (2, 337.0 – 372.0 mm SL); NMW 62940 (4, 191.5 – 235.0 mm SL); NMW 62947 (2, 337.0 – 372.0 mm SL); “ rio Doce, Steindachner Don., 1912 – 1913 ”.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F4EFFC64EA4F8C8FE87FCDF.taxon	diagnosis	Diagnosis. Brycon ferox can be diagnosed from all remaining cis-andean Brycon species, with the exception of B. stolzmanni, B. coxeyi, B. coquenani, B. vermelha, B. insignis, B. howesi, B. dulcis, B. vonoi, B. opalinus, and B. nattereri by possessing a color pattern consisting in a humeral blotch and a caudal peduncle blotch, without body stripes or other obvious color markings on caudal and anal-fins (vs. body stripes and caudal / anal fin color markings present; see Fig. 5). Brycon ferox can be distinguished from Brycon stolzmanni, B. coxeyi, B. vonoi, B. opalinus, and B. nattereri by possessing a distinctly acute head profile (vs. a roughly rounded to slightly acute head profile). Brycon ferox can be additionally distinguished from B. stolzmanni, and B. coxeyi by the absence of a patch of dark pigmentation on the opercle (vs. dark patch of pigmentation present on opercle). Brycon ferox can be distinguished from B. insignis, B. howesi, B. coquenani, and B. vermelha by possessing by possessing a fifth infraorbital bone about as wide as high (vs. fifth infraorbital bone wider than high; see Fig. 6). Brycon ferox can be distinguished from B. dulcis by presenting a distinctly anisognathous mouth, with a pointed premaxillary, extending beyond dentary, leaving the outer, and often also the second, series of premaxillary teeth exposed in ventral view (vs. premaxillary and dentary approximately isognathous, i. e., of about the same size, leaving only part of outer premaxillary series exposed in ventral view in some specimens). See the item “ Remarks ” of Brycon vonoi, for additional notes on the recognition of the species.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F4EFFC64EA4F8C8FE87FCDF.taxon	description	Description. Morphometric data are presented in Table 10. Large-sized species, largest examined specimen 375.0 mm SL. Body moderately slender in specimens up to 255 mm SL, moderately high in specimens> 310 mm SL. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly concave to straight 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 moderately acute anteriorly, considerably acute in specimens <100 mm SL (MZUSP 70219, 105.2 mm SL; MZUSP 93920, 2, 32.2 – 38.5 mm SL), mouth terminal. Jaws anisognathous, premaxillary moderately to pronouncedly projected relative to dentary, leaving outer row, and, in large specimens (e. g., MCZ 60934, 310.3 mm SL), also the second row of premaxillary teeth exposed in ventral view when mouth is closed. Maxillary long, extending posteriorly to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Six (1), 7 (5), 8 (6), 9 (11), 10 (10), or 11 (1) tricuspidate teeth in outer series. Two (1), 3 (2), 4 (11), 5 (16), 6 (2), or 7 (3) tetra- to hexacuspidate teeth in second, inner premaxillary row, plus 2 (14), 3 (19), or 4 (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, both pentacuspidate. Maxillary with distal portion distinctly expanded and rounded in profile. Seventeen to 31 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 9 (1), 10 (4), 11 (6), 12 (7), 13 (4), 14 (6), or 15 (1) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, penta- to heptacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, penta-, tetra-, tri- 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 8 (1), 14 (1), 17 (1), or 28 (1) small, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of sixth to ninth main series dentary teeth. Inner symphyseal teeth present in all specimens with relatively intact symphyseal dentary area. Central cusp distinctly larger and pointed, and teeth presenting diastemas in specimens <100 mm SL. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-eight (1), 49 (2), 51 (3), 52 (3), 53 (1), 54 (8), 55 (6), 56 (2), 57 (5), 58 (1), 60 (1), or 63 (1) scales in lateral line series. Laterosensory tube simple in small (<180 mm SL) specimens, deflected upwards in the first 5 – 6 scales, downwards in the remaining lateral-line scales. Larger specimens (> 190 mm SL) with branched tubules, mostly bifurcated but some scales with 3 tubules. Horizontal scale rows between dorsal-fin origin and lateral line eight (1), 9 (10), 10 (21), or 11 (2). Horizontal scale rows between lateral line and pelvic-fin 4 (8), 5 (22), or 6 (4). Circumpeduncular scales 15 (1), 16 (1), 17 (3), 18 (13), 19 (12), 20 (3), or 21 (1). Dorsal-fin rays ii, 9, a single specimen ii, 8. Dorsal fin origin slightly ahead of middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 13 th (1) vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 22 (1), 23 (2), 24 (7), 25 (9), 26 (10), or 27 (5). First anal-fin pterygiophore inserting behind haemal spine of 23 th (1) vertebra. Last unbranched and anterior 4 – 5 branched analfin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 25 – 30 per fin-ray main branch) middle-sized hooks on last unbranched and posterior main branch of branched rays 14 – 24, associated with dense, gelatinous tissue in 5 specimens (MZUSP 58048, 240.5 mm SL; MZUSP 70216, 185.3 mm SL; MNRJ 18379, 208.8 mm SL; NMW 62930, 1, 245.0 mm SL; NMW 62937, 1, 265.0 mm SL). A single hook per ray segment. Sheath of scales covering basis of anal-fin rays composed of two scale rows, lower scale row formed by 17 – 20 rectangular scales. Pectoral-fin rays i, 11 (1), 12 (4), 13 (19), or 14 (10). Pelvic-fin rays i, 7, a single specimen i, 6. Main caudal-fin rays 10 / 9. Caudal fin forked, lobes pointed. Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First branchial arch with 9 (1), 12 (4), 13 (5), 14 (5), or 15 (1) lower, 1 at angle, and 8 (1), 11 (3), 12 (6), 13 (4), or 14 (2) upper gill rakers. Vertebrae 44 (1). Supraneurals 10 (1). Coloration in alcohol. Overall body coloration clear. Top of head, snout, supraorbital, sixth infraorbital, upper portion of fifth infraorbital, and dorsal portion of body light-brown. Second, third, and fourth infraorbitals, and opercle silvery. Dentary, maxillary, gular area and lower portion of body light brown. Lateral portion of body cream-colored, with a silvery hue. Humeral blotch present, generally little 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 fourth to fifth lateral line scales, and vertically less than one scale high. Large, little to moderately conspicuous, oval-shaped caudal peduncle blotch, extending along 8 – 10 last lateral-line scales. All rayed fins with some amount of dark pigmentation at the interradial membranes, imparting an overall darkened coloration to these fins. Caudal-fin with a poorly discernible, roughly V-shaped blotch, formed by dark pigmentantion situated on outer caudal-fin rays. Adipose fin light-grey to light-brown. Color in life. Description based on the picture of a freshly collected (LIRP 1311, 310.0 mm SL; Lima & Castro, 2000: 157, fig. 2, lower specimen) and in a picture of a unpreserved specimen from the rio Mucuri (Silva, 1999: 16 – 17). Infraorbital bones, opercle and sides of body silvery. Opercle with some golden hue. Top of head, snout and dorsum dark-colored. Relatively conspicuous dark humeral and caudal peduncle blotches. Darkened dorsal, caudal, anal and pectoral fins. Specimen LIRP 1311 with series of pinkish spots situated along the four horizontal scale rows situated below scale row immediately below lateral line and anterior to insertion of pelvic fins. Specimens MZUSP 70215 (2, 184.4 – 229.4 mm SL) also displayed these pinkish spots when freshly preserved, in 2001. Sexual dimorphism. Steindachner (1877: 25 – 26) remarked that two of the syntypes were males and possessed small “ denticles ” (= “ Zähnchen ”) in the anal fin. We examined these two syntypes (NMW 62930, 1, 245.0 mm SL; NMW 62937, 1, 265.0 mm SL), plus eight specimens displaying anal-fin hooks (MZUSP 58048, 240.5 mm SL; MZUSP 70216, 185.3 mm SL; MNRJ 18379, 208.8 mm SL; CAS 13206, 5, 227.0 – 258.0 mm SL). Specimens MZUSP 58048, MZUSP 70216, and MNRJ 18379 were dissected and are males, with moderately well-developed testicles. Four specimens that did not displayed anal-fin hooks (MZUSP 70215, 182.8 mm SL; MZUSP 70216, 196.6 – 214.4 mm SL; MNRJ 18379, 255.6 mm SL) proved to be females, with moderately well-developed ovaries. Common names. “ Piabanha ” (Lima & Castro, 2000: 159).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F4EFFC64EA4F8C8FE87FCDF.taxon	distribution	Distribution. Known from coastal river drainages from northern Espírito Santo (rio Itaúnas) northward to southern Bahia (rio dos Frades drainage), including the rio Mucuri, rio Jucuruçu, and rio Itanhém drainages in the states of Bahia and Minas Gerais, eastern Brazil (Fig. 32). Ecological notes. Lima & Castro (2000: 161) remarked on the syntopy between Brycon ferox and Brycon vermelha at the middle rio Mucuri, Minas Gerais. However, intensive fish collecting at the rio Mucuri basin demonstrated that Brycon ferox is actually more common at the middle and lower stretches of that river system, being replaced by B. vermelha on the upper stretches (P. Pompeu and F. Vieira, pers. comm.). Brycon ferox is even reported for the lower stretch of the rio Mucuri, near the estuary (Lopes, 2000, F. Vieira and P. Pompeu, pers. comm.). Lima & Castro (2000: 161) found a whole Bolomys lasiurus (Rodentia, Cricetidae) in the stomach of Brycon ferox (LIRP 1311, 310.0 mm SL). The species obviously undertakes a spawning migration, as indicated by the great number of individuals moving upstream during the rainy season (November to March) (Pompeu & Martinez, 2006).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F4EFFC64EA4F8C8FE87FCDF.taxon	description	Conservation. Contrary to most Brycon species occurring at eastern Brazil, Brycon ferox currently does not appear to be endangered. The species was still quite common at the middle and lower rio Mucuri during the early 2000 ’ s (F. Vieira and P. Pompeu, pers. comm.), although populations of the species were probably adversely impacted after the building of the Santa Clara dam in 2002 and of the Mucuri dam in 2013. Aparently the species does not depend on riparian forests, since it is relatively common in deforested river stretches (F. Vieira and P. Pompeu, pers. comm.). The species was recently recorded for several small coastal river systems: the rio dos Frades, rio Itanhém, and rio Jucuruçu in southern Bahia and at the rio Itaúnas at northern Espírito Santo. These river systems possess watersheds almost entirely deforested, and the occurrence of Brycon ferox at those rivers apparently indicates some tolerance with this type of anthropogenic impact. However, small hydroelectric dams are being built in some of these rivers (e. g., at the rio Jucuruçu), and Brycon ferox populations will probably be adversely impacted.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F4EFFC64EA4F8C8FE87FCDF.taxon	discussion	Remarks. Steindachner (1877) described Brycon ferox from an unspecified locality at the rio Mucuri, eastern Brazil. The type material was evidently collected by the Thayer Expedition, since almost all material from eastern Brazil studied by Steindachner was brought by him from the MCZ (Steindachner, 1875: 499 – 500). As discussed under the item “ Remarks ” of Brycon insignis, we prefer not to consider specimens from MCZ as possessing type status except when Steindachner explicitly mention material deposited in that collection, which does not happen to be the case with B. ferox. There are three syntypes of Brycon ferox deposited at the NMW, the better preserved one (NMW 62930, 1, 245.0 mm SL) being here designated as the lectotype of the species, the two remaining specimens (NMW 62937, 247.0 – 265.0 mm SL) becoming, thus, paralectotypes. Interestingly, Howes (1982: 28) suggested that Brycon acuminatus (= B. insignis) might be a synonym of B. ferox. That supposition was based on the examination of the illustration of Brycon ferox by Steindachner (1877). Since the illustration indicated as being Brycon ferox in Steindachner (1877) actually represents B. insignis (Steindachner, 1879 c: 50, footnote), Howes (1982) was actually suggesting that B. acuminatus was probably a synonym of B. insignis, an assumption that was proved to be true in the present study (see section “ Remarks ” of Brycon insignis, above).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F4EFFC64EA4F8C8FE87FCDF.taxon	materials_examined	Material examined. Type material: NMW 62930 (1, 245.0 mm SL): “ Rio Mucuri ” “ I. 1874 ” [rio Mucuri basin, eastern Brazil; collected by C. F. Hartt and E. Copeland, 1865 – 1866; cf. Dick, 1977, Higuchi, 1996]. Lectotype (by present designation) of Brycon ferox. NMW 62937 (2, 247.0 – 265.0 mm SL): same data as lectotype; paralectotypes. Non types. Brazil, Minas Gerais: MZUSP 53305 (1, 210.4 mm SL); USNM 320346 (1, 203.7 mm SL); LIRP 1311 (1, 310.0 mm SL); Carlos Chagas, rio Mucuri, approx. 9 km W of Presidente Pena village, dirt road on Fazenda Gavião, c. 17 ° 37 ' S, 40 ° 55 ' W; R. M. C. Castro & S. L. Jewett, 17 – 23 Jul 1991. MZUSP 58048 (1, 240.5 mm SL), Carlos Chagas, rio Mucuri at Carlos Chagas, c. 17 ° 41 ’ S, 40 ° 46 ’ W; V. Vono, 14 Jan 1997. LBP 9065 (4, 133.9 – 235.0); LBP 10181 (5, 78.4 – 88.1 mm SL); LBP 9067 (4, 298.0 – 375.0 mm SL): Carlos Chagas, rio Mucuri, c. 17 ° 41 ’ S, 40 ° 46 ’ W; J. A. Senhorini, 6 Feb 2010. LBP 8099 (2, 223.0 – 268.0 mm SL); LBP 8100 (2, 92.9 – 97.4 mm SL): Carlos Chagas, rio Mucuri, 17 ° 41 ’ 42 ’’ S, 40 ° 46 ’ 11 ’’ W; C. Oliveira et al., 18 May 2009. MZUSP 70215 (8, 182.8 – 238.8 mm SL), Nanuque, rio Mucuri, downstream Tombo, c. 17 ° 51 ’ S, 40 ° 18 ’ W; F. Vieira, April 2001. Bahia: MCZ 60934 (1, 310.3 mm SL); MCZ 21108 (1, 375.0 mm SL); MCZ 21107 (3, 197.4 – 202.3 mm SL): CAS 13206 (5, 227.0 – 258.0 mm SL): rio Mucuri at Santa Clara; 17 ° 54 ' S, 40 ° 13 ' W; C. F. Hartt & E. Copeland, Dec 1865 — April 1866. USNM 301696 (1, 264.6 mm SL), rio Mucuri drainage, rio Mucuri approx. 26 km SE of town of Nanuque on Fazenda Santa Clara, 17 ° 54 ' S, 40 ° 13 ' W; S. L. Jewett & R. M. C. Castro, et al., 4 – 5 Aug 1988. MZUSP 70216 (4, 185.3 – 214.4 mm SL), Argolo, rio Mucuri, c. 17 ° 56 ’ S, 40 ° 7 ’ W; F. Vieira, April 2001. MZUSP 70219 (1, 105.2 mm SL), Mucuri, rio Mucuri, near its mouth, c. 18 ° 5 ’ S, 39 ° 34 ’ W; F. Vieira & P. S. Pompeu, Jan 2001. UFBA 5095 (2 of 5, 83.6 – 107.0 mm SL): Itamaraju, rio Jucuruçu (= Braço Sul), road BR- 101, 17 ° 14 ’ 52 ’’ S, 39 ° 37 ’ 15 ’’ W; A. M. Zanata et al., 28 Feb 2009. UFBA 5060 (1, 128.1 mm SL): Teixeira de Freitas, rio Itanhém, Prainha village, near road BR- 101, 17 ° 30 ’ 9 ’’ S, 39 ° 41 ’ 59 ’’ W; A. M. Zanata et al., 28 Feb 2009. MZUSP 93920 (2, 32.2 – 38.5 mm SL): Trancoso, rio dos Frades, 16 ° 38 ’ 39 ’’ S, 39 ° 8 ’ 32 ’’ W; N. A. Menezes, O. T. Oyakawa, J. C. Nolasco & L. M. Sousa, 1 April 2006. Espírito Santo: MNRJ 18379 (3, 192.3 – 255.6 mm SL), Pedro Canário, rio Itaúnas, Fazenda Boa Lembrança, c. 18 ° 19 ’ S, 39 ° 50 ’ W; A. C. Aguirre, 1950. UFBA 5021 (2 of 3, 87.4 – 115.6 mm SL): Pedro Canário, rio da Samambaia (trib. rio Itaúnas), road BR- 101, 18 ° 12 ’ 24 ’’ S, 39 ° 55 ’ 56 ’’ W; A. M. Zanata et al., 27 Feb 2009.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F49FFB94EA4FC4BFF04F8AA.taxon	diagnosis	Diagnosis. Brycon vonoi can be diagnosed from all remaining cis-andean Brycon species, with the exception of B. stolzmanni, B. coxeyi, B. coquenani, B. vermelha, B. insignis, B. howesi, B. dulcis, B. ferox, B. opalinus, and B. nattereri by possessing a color pattern consisting in a humeral blotch and a caudal peduncle blotch, without body stripes or other obvious color markings on caudal and anal-fins (vs. body stripes and caudal / anal fin color markings present; see Fig. 5). Brycon vonoi can be distinguished from B. ferox, B. vermelha, B. insignis, B. howesi, B. coquenani, and B. dulcis by possessing a slightly acute head profile (vs. a distinctly acute head profile). Brycon vonoi can be distinguished from Brycon stolzmanni and B. coxeyi by the absence of a patch of dark pigmentation on the opercle (vs. dark patch of pigmentation present on opercle). Brycon vonoi can be additionally distinguished from B. insignis, B. howesi, B. coquenani, and B. vermelha by possessing a fifth infraorbital bone about as wide as high (vs. fifth infraorbital bone wider than high; see Fig. 6). Brycon vonoi can be additionally distinguished from B. dulcis by possessing a clear overall color pattern (vs. darkened overall body color). Brycon vonoi can be distinguished from B. nattereri by possessing lateral line tubules mostly simple (vs. tubules with 2 – 5 branches), caudal peduncle blotch not extending into middle caudal-fin rays (vs. caudal peduncle blotch extending into middle caudal-fin rays), and roughly V-shaped blotch on outer caudal-fin rays present (vs. outer caudal-fin rays clear, without dark pigmentantion). Brycon vonoi can be distinguished from B. opalinus by presenting an acute, pointed snout (vs. obtuse, rounded snout) and a longer upper jaw length (45.1 – 51.4 % of HL, mean 49.5, vs. 38.7 – 48.3 of HL, mean 42.9).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F49FFB94EA4FC4BFF04F8AA.taxon	description	Description. Morphometric data are presented in Table 11. Middle-sized species, largest examined specimen 278.0 mm SL. Body moderately slender. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly concave 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 isognathous to slightly anisognathous, premaxillary projecting slightly relative to dentary in some specimens, leaving outer row of premaxillary teeth exposed when mouth is closed. Maxillary long, extending posteriorly to anterior third to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Five (1), 6 (8), 7 (7), 8 (2), 9 (6), 10 (1), or 11 (2) tricuspidate teeth in outer series. Three (3), 4 (11), 5 (10), or 6 (2) tri- to tetracuspidate teeth in second, inner premaxillary row, plus 2 (2), 3 (12) or 4 (12) 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, both pentacuspidate. Maxillary with distal portion distinctly expanded and rounded in profile. Seventeen to 27 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 9 (2), 10 (2), 11 (3), 12 (1), or 14 (2) 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, tetra-, tri- 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 25 – 31 small, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of third main series dentary teeth. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-five (3), 46 (4), 47 (13), 48 (6), or 49 (2) scales in lateral line series. Laterosensory tube generally simple, deflected upwards in the first 5 – 6 scales, downwards in the remaining lateral-line scales. Larger specimens (> 230 mm SL) with with tubules bifurcated in some scales. Horizontal scale rows between dorsal-fin origin and lateral line 6 (1), 7 (16) or 8 (11). Horizontal scale rows between lateral line and pelvic-fin 4 (22) or 5 (6). Circumpeduncular scales 14 (1), 15 (2), 16 (14), 17 (9), or 19 (1). Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead of middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 13 th (1) or 14 th (1) vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 20 (1), 21 (7), 22 (12), 23 (4), or 25 (3). First anal-fin pterygiophore inserting behind haemal spine of 23 th (2) vertebra. Last unbranched and anterior 4 – 5 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 25 – 30 per fin-ray main branch) middle-sized hooks on last unbranched and posterior main branch of branched rays 11 – 23, associated with dense, gelatinous tissue in 7 specimens (MZUSP 58297, 1, 210.6 mm SL; MZUSP 93815, 2, 195.2 – 192.3 mm SL; ZUEC 10746, 1, 189.6 mm SL; MCP 49962, 1, 194.1 mm SL; MCNIP 1595, 1, 193.4 mm SL; MCNIP 1596, 1, 182.6 mm SL), one of which (MZUSP 58297) a female (see “ Sexual dimorphism ”, below). A single hook per ray segment. Sheath of scales covering basis of anal-fin rays composed of two scale rows, lower scale row formed by 20 – 23 rectangular scales. Pectoral-fin rays i, 10 (2), 11 (8), or 12 (17). Pelvic-fin rays typically i, 7, one specimen i, 8. Main caudal-fin rays 10 / 9. Caudal fin forked, lobes rounded. Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First branchial arch with 11 (1), 13 (1), or 16 (1) lower, 1 at angle, and 11 (1), 12 (1), or 13 (1) upper gill rakers. Vertebrae 43 (2). Supraneurals 10 (1) or 11 (1). Coloration in alcohol. Overall body coloration clear. Top of head, snout, supraorbital, sixth infraorbital, and dorsal portion of body grayish to brownish. Second, third, fourth, and fifth infraorbitals, and opercle silvery. Dentary, maxillary, gular area and lower portion of body light brown. Lateral portion of body clear, with a silvery hue. Humeral blotch present, moderately conspicuous, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of second, extending longitudinally to posterior margin of fourth lateral line scales, and vertically less than one scale high. Large, little to moderately conspicuous, oval-shaped caudal peduncle blotch, extending along 5 – 6 last lateral-line scales. Rayed fins clear, with dark chromatophores scattered over the interradial membranes. Caudal-fin with a poorly discernible, roughly V-shaped blotch, formed by dark pigmentantion situated on outer caudal-fin rays. Adipose fin light-grey to light-brown. Color in life. The holotype (MZUSP 58297), a year and a half after being collected (September 1999), still retained orange blotches at the centre of the scales from the two horizontal scale rows situated below lateral line scale row, anterior to insertion of pelvic fins. Sexual dimorphism. Seven specimens (MZUSP 58297, 1, 210.6 mm SL; MZUSP 93815, 2, 195.2 – 192.3 mm SL; ZUEC 10746, 1, 189.6 mm SL; MCP 49962, 1, 194.1 mm SL; MCNIP 1595, 1, 193.4 mm SL; MCNIP 1596, 1, 182.6 mm SL) displayed anal-fin hooks. One of these specimens (MZUSP 93815, 192.3 mm SL) was dissected and is a male, with well-developed testicles. On other hand, the specimen MZUSP 58297 is a female, presenting oocytes at the urogenital opening. Another obviously mature female specimen, presenting oocytes at the urogenital opening (MZUSP 93815, 278.0 mm SL) does not present fin hooks. Fin hooks are generally regarded as being a dimorphic feature, present only in males (Malabarba & Weitzman, 2003). In fact, data for other Brycon species examined in the present study confirm this view. Consequently, the occurrence of anal-fin hooks in the mature female specimen MZUSP 58297 should be considered an abnormal condition.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F49FFB94EA4FC4BFF04F8AA.taxon	etymology	Etymology. The specific name honors our dear colleague and friend Volney Vono (1961 – 2011), who was the first ichthyologist to collect the new species and who brought it to the attention of the author. Common names. “ Piabanha ” (V. Vono, O. T. Oyakawa, pers. comm.).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F49FFB94EA4FC4BFF04F8AA.taxon	distribution	Distribution. Only known, and apparently endemic from the rio Pardo basin, a small coastal drainage situated at Minas Gerais and Bahia states, immediately north from the rio Jequitinhonha basin, eastern Brazil (Fig. 32). Conservation. The original Atlantic forest which formerly covered most of the rio Pardo basin was historically largely replaced by pastureland and crops. There is a hydroelectric dam (Machado Mineiro dam) build in the middle rio Pardo that very likely has affected adversely the species. Unfortunately, more hydroelectric dams are being planned for the rio Pardo basin. Brycon vonoi is being artificially bred at a fish hatchery situated at the Machado Mineiro hydroelectric dam. Probably, as almost all remaining eastern brazilian Brycon species, the species is threatened with extinction, though it is necessary to obtain additional field data to ascertain its current conservation status.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F49FFB94EA4FC4BFF04F8AA.taxon	discussion	Remarks. There is an old lot (MCZ 2416) said to have been collected at the “ rio Una ” by “ A. de Lacerda. Antônio de Lacerda was an amateur brazilian naturalist resident at Salvador, Bahia, who assisted N. Dexter and S. V. R. Thayer from the Thayer Expedition during their stay at that city (Dick, 1977: 6; Agassiz & Agassiz, 1975: 93). The lot MCZ 2416 was collected previously (1864) to the Thayer Expedition, presumably at the rio Una which lies at the city of Valença, where Lacerda’s family owned a cottom mill industry. This locality lies considerably to the north (~ 220 km) from the mouth of the rio Pardo, the type-locality of Brycon vonoi. These specimens are similar to Brycon vonoi and are herein provisionally considered to be conspecific to this species. More collecting in the coastal rivers of central and northern Bahia are, however, necessary to understand the taxonomical status of Brycon populations in the area (see also “ Remarks ” of Brycon devillei and B. opalinus).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F49FFB94EA4FC4BFF04F8AA.taxon	materials_examined	Material examined. Holotype: MZUSP 58297 (1, 210.6 mm SL): Brazil, Minas Gerais, Taiobeiras, rio Pardo, Fazenda Tabatinga, c. 15 ° 43 ’ S 42 ° 14 ’ W; V. Vono, 17 Feb 1998. Paratypes. Brazil, Minas Gerais, rio Pardo basin. MZUSP 104020 (1, 151.0 mm SL): same data as holotype. MZUSP 83434 (3, 149.2 – 232.0 mm SL): Berizal, rio Pardo, c. 15 ° 35 ' S, 41 ° 44 ' W; B. P. Nogueira & M. F. G. Brito, 23 Jun 2000. MCNIP 1595 (2, 178.6 – 193.4 mm SL); ZUEC 10746 (2, 166.0 – 189.6 mm SL); ANSP 200241 (1, 165.0 mm SL); MCP 49962 (1, 194.1 mm SL); MNRJ 45957 (1, 164.0 mm SL): Águas Vemelhas, rio Pardo, immediatelly below Machado Mineiro hydroeletric dam, 15 ° 31 ' 25 '' S, 41 ° 30 ' 26 '' W; T. C. Pessali et al., 15 Feb 2015. MZUSP 93815 (17, 82.1 – 278.0 mm SL, 2 cs, 84.9 – 92.8 mm SL): Águas Vermelhas, CEMIG fish hatchery (stocked from specimens collected at the rio Pardo), UHE Machado Mineiro, 15 ° 31 ’ 19 ’’ S, 41 ° 30 ’ 18 ’’ W; O. T. Oyakawa, J. L. Birindelli & L. M. Sousa, 16 Apr 2007. MZUSP 108439 (1, 124.6 mm SL): Ninheira, Rio Pardo, UHE Machado Mineiro, 15 ° 31 ’ S, 41 ° 31 ’ W; W. A. Meireles, March 2011. MCNIP 1596 (11, 103.0 – 182.6 mm SL); ZUEC 10747 (4, 105.1 – 113.7 mm SL): Ninheira, CEMIG fish hatchery (stocked from specimens collected at the rio Pardo), UHE Machado Mineiro, 15 ° 31 ’ 19 ’’ S, 41 ° 30 ’ 18 ’’ W; T. C. Pessali et al., 14 Feb 2015. Additional material (not typical). UFMG uncat. (1, 221.6 mm SL): same data as holotype. MCZ 2416 (2, 200.0 – 260.5 mm SL): Bahia, “ Rio Una ” [c. 13 ° 22 ’ S, 39 ° 5 ’ W; see Remarks, above]; A. de Lacerda, 18 March 1864.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F35FFB24EA4FF50FB06FC2F.taxon	description	Brycon cf. reinhardti (not Lütken): Narahara, 1993: 5 – 6 (gonadal development, maturation in captivity); Almeida-Toledo et al., 1996: 36 – 37 (karyotype).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F35FFB24EA4FF50FB06FC2F.taxon	diagnosis	Diagnosis: Brycon opalinus can be diagnosed from all remaining cis-andean Brycon species, with the exception of B. stolzmanni, B. coxeyi, B. coquenani, B. vermelha, B. insignis, B. howesi, B. dulcis, B. ferox, B. vonoi, and B. nattereri by possessing a color pattern consisting in a humeral blotch and a caudal peduncle blotch, without body stripes or other obvious color markings on caudal and anal-fins (vs. body stripes and caudal / anal fin color markings present; see Fig. 5). Brycon opalinus can be distinguished from Brycon ferox, B. vermelha, B. insignis, B. howesi, B. coquenani, and B. dulcis by possessing a rounded, obtuse head profile (vs. a distinctly acute head profile). Brycon opalinus can be distinguished from Brycon stolzmanni and B. coxeyi by the absence of a patch of dark pigmentation on the opercle (vs. dark patch of pigmentation present on opercle). Brycon opalinus can be additionally distinguished from B. insignis, B. howesi, B. coquenani, and B. vermelha by possessing a fifth infraorbital bone about as wide as high (vs. fifth infraorbital bone wider than high; see Fig. 6). Brycon opalinus can be distinguished from B. nattereri by possessing lateral line tubules mostly simple (vs. tubules with 2 – 5 branches), and caudal peduncle blotch not extending into middle caudal-fin rays (vs. caudal peduncle blotch extending into middle caual-fin rays). Brycon opalinus can be distinguished from B. vonoi by presenting an obtuse, rounded snout (vs. acute, pointed snout) and a shorter upper jaw length (38.7 – 48.3 % of HL, mean 42.9, vs. 45.1 – 51.4 of HL, mean 49.5).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F35FFB24EA4FF50FB06FC2F.taxon	description	Description. Morphometric data are presented in Table 12. Middle-sized species, largest examined specimen 278.0 mm SL. Body slender to moderately slender. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly concave 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 isognathous to slightly anisognathous, premaxillary projecting slightly relative to dentary in some specimens, leaving outer row of premaxillary teeth exposed when mouth is closed. Maxillary moderately long, extending posteriorly to anterior margin to anterior third of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Five (3), 6 (17), 7 (32), 8 (23), 9 (11), or 10 (1) tricuspidate teeth in outer series. Three (24), 4 (53), or 5 (7) tri- to tetracuspidate teeth in second, inner premaxillary row, plus 2 (16), 3 (53), or 4 (17) 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, both pentacuspidate. Maxillary with distal portion distinctly expanded and rounded in profile. Ten to 22 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with seven (3), 8 (12), 9 (20), 10 (15), 11 (9), 12 (4), or 13 (2) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, tetra-, tri- 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 – 16 small, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of fourth to fifth main series dentary teeth. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-two (1), 44 (1), 45 (2), 46 (9), 47 (3), 48 (22), 49 (15), 50 (15), 51 (5), 52 (9), 53 (3), 54 (2), 57 (1), 58 (1), or 59 (2) scales in lateral line series. Laterosensory tube generally simple, deflected upwards in the first 5 – 6 scales, downwards in the remaining lateral-line scales. Some large specimens (> 230 mm SL) with tubules bifurcated in some scales. Horizontal scale rows between dorsal-fin origin and lateral line seven (3), 8 (56), 9 (30), or 10 (4). Horizontal scale rows between lateral line and pelvic-fin 4 (55), 5 (36), or 6 (1). Circumpeduncular scales 14 (1), 15 (14), 16 (43), 17 (19), 18 (12), or 19 (4) Dorsal-fin rays typically ii, 9, two specimens ii, 10. Dorsal fin origin at middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 14 th (1) or 15 th (3) vertebrae. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 17 (2), 18 (7), 19 (24), 20 (39), 21 (18), 22 (10), or 23 (2). First anal-fin pterygiophore inserting behind haemal spine of 25 th (2), 26 th (1), or 28 th (1) vertebrae. Last unbranched and anterior 4 – 5 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 15 – 22 per fin-ray main branch) small hooks on last unbranched and posterior main branch of branched rays 16 – 22, associated with dense, gelatinous tissue in two specimens (MZUSP 50805, 224.6 mm SL; MZUSP 59280, 215.5 mm SL). A single hook per ray segment. Sheath of scales covering basis of anal-fin rays composed of two scale rows, lower scale row formed by 13 – 20 rectangular scales. Pectoral-fin rays i, 11 (10), 12 (38), 13 (43), or 14 (2). Pelvic-fin rays typically i, 7, two specimens i, 6. One specimen (MZUSP 50805, 224.6 mm SL) with minute hooks on distal portion of posterior branch of branched pelvic-fin rays. Main caudal-fin rays 10 / 9. Caudal fin forked, lobes rounded. Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First branchial arch with 10 (6), 11 (10), 12 (20), 13 (19), or 14 (3) lower, 1 at angle, and 8 (1), 9 (16), 10 (21), 11 (15), or 12 (3) upper gill rakers. Vertebrae 43 (1), 45 (1), 46 (2), or 47 (2). Supraneurals 10 (2), 11 (1), or 12 (1). Coloration in alcohol. Overall body coloration brown. Top of head, snout, supraorbital, fifth and sixth infraorbitals, and dorsal portion of body light- to dark-brown. Second, third, fourth, and fifth infraorbitals, and opercle light-brown, silvery in specimens retaining guanine. Dentary, maxillary, gular area and lower portion of body light brown. Lateral portion of body light-brown, with a silvery hue in specimens retaining guanine. Humeral blotch present, relatively conspicuous in juveniles but little conspicuous in adult specimens, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of second, extending longitudinally to posterior margin of fourth lateral line scales, and vertically one and half scales high. Large, little conspicuous, oval-shaped caudal peduncle blotch, extending along 6 – 7 last lateral-line scales. Rayed fins clear, with dark chromatophores scattered over the interradial membranes. Caudal-fin with a poorly discernible, roughly V-shaped blotch, formed by dark pigmentantion situated on outer caudal-fin rays in some specimens. Adipose fin light-grey to light-brown. Specimens from rio Santo Antônio basin in upper rio Doce drainage with overall body color, including fins, darkened, possibly a result of the dark waters of that river system. Color in life. Description based on pictures of unpreserved specimens from a cultivated stock kept at the CESP-Paraibuna fish hatchery and in specimens collected at the rio Paraibuna (MZUSP 51538). Dorsal area, top of head and snout darkened. Infraorbitals and opercular bones silvery, with a diffuse dark pigmentantion. Lateral portion of body clear, with a silvery hue. Conspicuous dark peduncular blotch. Pectorals, pelvics, anal and caudal fins darkened. Sexual dimorphism. Two specimens (MZUSP 50805, 224.6 mm SL; MZUSP 59280, 215.5 mm SL) with small hooks present on posterior branch of branched anal- and (MZUSP 50805 only) pelvic-fin rays. Both specimens were dissected and proved to be mature males, with very well-developed testes. An obviously reproductive female specimen (MZUSP 59282, 249.5 mm SL), with oocytes on the urogenital opening, lacks fin hooks. Common names. “ Pirapitinga-do-sul ”; “ pirapitinga ”; “ pipitinga ” (MG); “ parpitinga ” (SP).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F35FFB24EA4FF50FB06FC2F.taxon	distribution	Distribution. Restricted to headwater tributaries from the rio Paraíba do Sul and rio Doce basins, eastern Brazil (Fig. 43). A old record for the lower rio Doce basin in Espírito Santo state (MCZ 21104) is here considered doubtful in view of the known preference of the species for headwater areas. See the item “ Remarks ”, below, for an account on the locality “ Bahia ” (type-locality of the holotype of Brycon bahiensis).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F35FFB24EA4FF50FB06FC2F.taxon	description	Variation. Specimens herein identified as Brycon opalinus generally agree in most morphometric and meristic characters. There is, however, slight morphological variation between different populations of the species. Specimens from the upper rio Paraibuna and rio Itaguaçaba at the upper rio Paraíba do Sul basin in São Paulo state and from rio Preto at the middle rio Paraíba do Sul basin in Rio de Janeiro / Minas Gerais states present a relatively lower body depth (21.3 – 29.8 % of SL) when compared with specimens from the rio do Peixe in the middle rio Paraíba do Sul basin in Minas Gerais (27.5 – 32.8 % of SL) and from specimens from the rio Santo Antônio basin (upper rio Doce drainage) in Minas Gerais (26.3 – 30.7 % of SL). The single cleared and stained Brycon opalinus from the rio Doce basin presented a lower total vertebral count compared with the four cleared and stained specimens from the rio Paraíba do Sul basin (43, vs. 45 – 47, respectively). Also, specimens from the upper rio Doce basin basin are generally darker than specimens from the rio Paraíba do Sul basin, though this difference is probably merely a phenotypic variation associated with the black waters where the Brycon opalinus population from the Rio Doce basin dwell (see color variation of B. howesi, B. dulcis, and B. nattereri for similar instances of dark and clear-colored populations). Since there is considerable overlap in morphometric data and no clear geographical patterns in the variation observed, we consider that all these populations should be considered a single species. Analyses of the variability of mitochondrial DNA (Hilsdorf et al., 2002) and microsatellite loci (Barroso et al., 2005) indicated some genetic variation among distinct Brycon opalinus populations from the upper and middle rio Paraíba do Sul basin which are, however, within the variability expected in a species with a low genetic interchange. Ecological notes. Brycon opalinus is exclusively found in headwater areas, inhabiting small rivers, characterized by possessing slightly dark-stained to clear waters, and lentic stretches alternated with rapids, as the rio Paraibuna in the upper rio Paraíba do Sul basin, the rio Preto at the middle portion of the same river system, and the rio Santo Antônio in the upper rio Doce basin. The diet of the species in the upper rio Paraibuna basin was studied by Gomiero et al. (2008), based on the examination of 256 Brycon opalinus stomach contents collected along a whole year. Brycon opalinus is an omnivore, ingesting animal and vegetal food items in about equal proportions, except for the juveniles, that show a tendency to ingest a greater proportion of animal food itens (Gomiero et al., 2008). Insects (mainly allochthnous), fruits and seeds were the main food itens found in stomach contents (Gomiero et al., 2008). An exceptional occurrence of frogs and a rat in stomach contents of Brycon opalinus in the upper rio Paraibuna was ascribed to have been caused by a rainstorm, which probably washed these animals into the river (Gomiero et al., 2006). Stomach contents of two examined specimens (MZUSP 59279, 195.9 mm SL; MZUSP 59280, 247.1 mm SL) from rio Santo Antônio basin was composed by crushed fruits and seeds and a beetle. Specimens collected at the rio Preto, presented stomach contents filled with larvae of Corydalidae (Megaloptera, Insecta; A. Hilsdorf, pers. comm.). Reproduction of Brycon opalinus was studied by Gomiero & Braga (2007 a, b) in the upper rio Paraibuna basin. As other Brycon species, B. opalinus is a total spawner, presenting two reproductive peaks, one during the spring / summer and other during the autumn (Gomiero & Braga, 2007 a). Individuals in breeding condition collected in other river basins confirm that reproduction occurs both during the summer and autumn, as evidenced by a mature male, with well-developed testicles and anal-fin hooks, colected at the rio do Peixe (middle rio Paraíba do Sul basin) during end of April and early May (MZUSP 50805), and a mature female, with oocytes at the urogenital opening, collected at the rio Preto do Itambé (upper rio Doce basin) in March (MZUSP 59282). Sexual maturity is achieved at its earliest with 16 cm TL, and mean fecundity was estimated as 9190 oocytes (Gomiero & Braga, 2007 a). Females grow larger than males, and the sex-ratio is skewed towards females (Gomiero & Braga, 2007 a, b). Specimens kept in captivity achieved the sexual maturity at the second year of age (Narahara, 1993). It is evident that, contrary to some of its congeners, Brycon opalinus does not perform long-distance reproductive migrations, though local migrations might occur in the upper rio Paraibuna (J. P. Villani, pers. comm., 1996). Similarly to some centro-american Brycon species (Kramer, 1978; Bussing, 1993), spawning in the upper rio Paraibuna occurs in small streams (J. P. Villani fide Gomiero & Braga, 2007 b). Growth and mortality parameters of Brycon opalinus in the upper rio Paraibuna basin were estimated by Gomiero et al. (2007) using total-length size classes frequencies. These authors inferred that growth slows during late winter / early spring and that longevity for males and females was 4.50 years. Those growth and aging estimates need to be however validated using more empirical methods such as analyses of ring growths of otoliths and scales (see Pauly, 1987, for a critique on statistical aging methods). Conservation. Brycon opalinus was presumably originally widely distributed across the rio Paraíba do Sul and rio Doce headwaters areas. Its current geographical range is highly fragmented and restricted to some headwater tributaries of the rio Paraíba do Sul and rio Doce basins. A healthy population of the species occurs within the limits of the Santa Virgínia unit of the Serra do Mar State Park. There are some other healthy populations of the species occurring in some areas, such as the ones occurring at the rio Preto basin in the middle rio Paraíba do Sul basin and at the rio Santo Antônio in the upper rio Doce basin, which unfortunately lack formal protection and are threatened by several ongoing anthropogenic activities, specially hydroelectric dams building. Stocking and reintroduction programs using population genetics information are under way for the species (Hilsdorf et al., 2002; Barroso et al., 2005). The species is formally considered as threatened with extinction in Brazil, under the category “ Vulnerable ” (see Hilsdorf, Lima & Vieira, 2008 for additional information). The recent catastrophic burst of a tailings dam in the upper rio Doce in November 2015, which released millions of tons of iron waste into the rio Doce basin and devastated his entire course, probably had no impact on the Brycon opalinus populations of this basin since they are restricted to some tributaries (rio Santo Antônio and rio Piranga) which were not directly affected by the iron waste.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F35FFB24EA4FF50FB06FC2F.taxon	discussion	Remarks. Cuvier (1819: 351 – 352) described Chalceus opalinus based on a specimen collected by the botanist Auguste de Saint-Hilaire at an unspecified locality in Brazil. Valenciennes (in Cuvier & Valenciennes, 1850: 244 – 246) redescribed a species based on two specimens, one collected at the “ Rio Tiquilenhonha ” (= rio Jequitinhonha), which was also collected by Saint-Hilaire, and another specimen from “ Rio Janeiro ”, collected by “ M. de Lalande ”. Although Valenciennes (in Cuvier & Valenciennes, 1850) based his description of Chalceus opalinus on the specimen from the rio Jequitinhonha, this specimen certainly does not correspond to the specimen studied by Cuvier (1819) because it was donated to the MNHN in August 1822 (Valenciennes, in Cuvier & Valenciennes, 1850: 246). In fact, the specimen collected by “ de Lalande ”, MNHN A. 8613, was, as previously remarked by Bertin (1948) and Géry & Mahnert (1992: 813) the one studied by Cuvier (1819), being, thus, the holotype of Chalceus opalinus. Pierre Antoine Delalande travelled with Saint-Hilaire to Brazil and accompanied him from Rio de Janeiro city to “ Ubá ”, a port at the banks of the rio Paraíba do Sul, and then returned to the Rio de Janeiro city and from there to France, carrying the collections obtained during this travel (Papavero, 1971). This probably explains why Cuvier (1819) mentioned Saint-Hilaire, instead of Delalande, as being the collector of the holotype of Chalceus opalinus. As earlier remarked by Eigenmann (1921: 294 – 295), the type locality of Chalceus opalinus should thus be restricted to “ Rio de Janeiro ”. It is thus very likely that the precise type-locality of Brycon opalinus is the rio Paraíba do Sul at “ Ubá ” (currently Andrade Pinto, 22 ° 14 ’ S, 43 ° 25 ’ W) where Saint-Hilaire and Delalande spent some time collecting insects (Papavero, 1971: 116). As for the specimen MNHN A. 8612, collected in the rio Jequitinhonha, it undoubtedly represents Brycon howesi (see item “ Remarks ” of this species, above). Günther (1864: 334) described Brycon bahiensis bearing the vague type-locality “ Bahia ”. Localities known as have been sampled by the collector of the holotype of Brycon bahiensis, Otho Wucherer, lies in the neighborings of Salvador (the capital of Bahia state) and some coastal localities southward from it (Nazaré, Canavieiras, and Ilhéus) (Günther, 1861, 1863; Wucherer, 1861 a, b; 1863 a, b). The comparison of the holotype of the species (BMNH 1862.11.23: 26) with Brycon opalinus specimens from the rio Paraíba do Sul and rio Doce basins did not revealed any significative differences between them. Howes (1982: 11) remarked that the holotype of Brycon bahiensis possess more premaxillary and maxillary teeth than a Brycon opalinus specimen from Porto Real, Rio de Janeiro (BMNH 1912.11.21: 1). Examination of a large sample of Brycon opalinus in the present study demonstrated that premaxillary counts of the holotype of B. bahiensis lies within the variation of B. opalinus, while the maxillary teeth number is only slightly above the range of variation found in B. opalinus (24, vs. 10 – 22, respectively). In spite of the fact that the presumable type-locality of Brycon bahiensis lies well outside the known geographical range of Brycon opalinus, we herein consider the former a junior synonym of the latter. Coastal river systems between Salvador and the rio Jequitinhonha mouth, which is the presumable type-locality area of Brycon bahiensis, are still poorly known and more collecting is necessary to ascertain the occurrence of B. opalinus, or a species close to it, in the area (see also the item “ Remarks ” of B. devillei and B. vonoi). At this time we have no better option left other than to consider the type-locality of Brycon bahiensis as probably incorrect, and that somehow its type specimen was collected southwards, within the geographical range of B. opalinus. Magalhães (1931: 168) recorded Brycon pirapitinga in São Luís do Paraitinga, a locality lying at the upper rio Paraíba do Sul basin. However, the name Salmo pira-pitinga Kner (ex Natterer) (1860) is invalid, since it was originally proposed as a synonym of Chalceus opalinus. Both the common name and the locality mentioned by Magalhães (1931) leaves no doubt that his record refers to Brycon opalinus. Several authors, starting with Steindachner (1877: 27 – 29), incorrectly identified specimens of Brycon opalinus from the rio Paraíba do Sul and rio Doce basins as Brycon reinhardti (which is a synonym of B. nattereri, see the item “ Remarks ” of this latter species).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F35FFB24EA4FF50FB06FC2F.taxon	materials_examined	Material examined. Type material. MNHN A. 8613 (1, 278 mm SL): “ Brésil ” [corrected to Rio de Janeiro by Eigenmann, 1921; see “ Remarks ”, above, for the probable precise locality]; P. A. Delalande & A. Saint-Hillaire, 1816. Holotype of Chalceus opalinus Cuvier. BMNH 1862.11.23: 26 (1, 128.2 mm SL): “ Bahia ”; O. Wucherer. Holotype of Brycon bahiensis Günther, 1864. Additional material. Brazil, São Paulo, rio Paraíba do Sul basin: MZUSP 21872 (12, 68.9 – 213.4 mm SL): rio Paraibuna, Alto da Serra; CESP personnel, May 1982. MZUSP 42059 (8, 84.4 – 239.0 mm SL): Paraibuna, Represa de Paraibuna; CESP personnel, no date. MZUSP 28338 (9, 146.9 – 181.4 mm SL): Paraibuna, CESP fish hatchery; CESP personnel, 16 Feb 1984. MZUSP 51361 (2, 130.0 – 170.2 mm SL): São Luiz do Paraitinga, rio Paraibuna, Salto Grande, 23 ° 20 ’ S, 45 ° 9 ’ W; CESP personnel, no date. MZUSP 46783 (1, 94.3 mm SL): São Luiz do Paraitinga, rio Turvo (trib. rio Paraibuna); R. S. Lima & I. C. Possagnolo, 26 Jul 1992. MZUSP 51358 (5, 131.4 – 158.6 mm SL): São Luiz do Paraitinga, rio Paraibuna, Salto Grande, Santa Virgínia, Serra do Mar State Park, 23 ° 20 ’ S, 45 ° 9 ’ W; F. C. T. Lima et al., 2 – 3 Nov 1996. MZUSP 19127 (5, 135.4 – 233.2 mm SL): Silveiras, rio Itaguaçaba basin, fazenda Marrano, c. 22 ° 39 ’ S, 44 ° 51 ’ W; Jabes T. Braga, May 1981. LBP 6303 (1, 148.8 mm SL): Areias, rio Itagaçaba, 22 ° 39 ’ 26 ’’ S, 44 ° 45 ’ 50 ’’ W; R. Devidé et al., 1 Apr 2008. Rio de Janeiro, rio Paraíba do Sul basin: MZUSP 16624 (1, 133.7 mm SL): Resende, rio Paraíba do Sul; P. Azevedo, 1962. MNRJ 11273 (1, 103.3 mm SL): rio Preto (bacia do Paraíba do Sul), Bagagem, Bocaina de Minas and Resende border, 22 ° 15 ' S, 44 ° 20 ' W; G. W. Nunan & W. D. Bandeira, 19 Sept 1982. MNRJ 11255 (5, 97.1 – 188.9 mm SL): rio Preto (bacia do rio Paraíba do Sul), 1.5 km downstream Resende-Bocaina de Minas road, borderline RJ / MG, c. 22 ° 15 ' S, 44 ° 20 ' W; J. R. S. Araújo, 6 Sept 1982. MNRJ 11255 (12, 75.3 – 208.9 mm SL): Resende, rio Preto, Bagagem-Fumaça road, c. 2 km from road Resende / Bocaina de Minas, c. 22 ° 15 ' S, 44 ° 20 ' W; J. R. S. Araújo, 22 Aug 1982. MNRJ 11256 (2. 93.5 – 160.0 mm SL): Resende, rio Pirapitinga (bacia do rio Paraíba do Sul), 3 – 4 km above Fazenda Monte Alegre; J. R. S. Araújo, 6 – 7 Sept 1982. MZUSP 83433 (2, 205.0 – 272.0 mm SL): Rio das Flores, rio Preto, c. 22 ° 5 ’ S, 43 ° 33 ’ W; B. P. Nogueira & G. V. Padilha, 19 Jul 2000. BMNH 1912.11.21: 1 (1, 155.4 mm SL): Porto Real, 22 ° 25 ’ S, 44 ° 17 ’ W; M. Hardy de Dréneuf. MNRJ 12227 (1, 94.3 mm SL): Teresópolis, rio Preto, at the confluence with rio Paquequer; D. F. Moraes Jr, 1980. MNRJ 15291 (1, 45.5 mm SL): Carmo, rio Paquequer (trib. left margin rio Paraíba do Sul), c. 1.5 km above bridge on road RJ- 160, 21 ° 53 ’ S, 42 ° 38 ’ W; D. F. Moraes Jr. et al., Sept 1989. MNRJ 6305 (1, 168.5 mm SL): Sapucaia, rio Paraíba [do Sul], 21 ° 56 ' S, 42 ° 20 ' W; C. Moreira, Feb 1911. MNRJ 21469 (1, 102.9 mm SL): Volta Grande, rio Paraíba do Sul, Ilha dos Pombos, 21 ° 50 ’ 16 ’’ S, 42 ° 34 ’ 56 ’’ W; W. A. Gosline, 12 – 14 Jan 1945. MCZ 21123 (1, 210.5 mm SL): Rio de Janeiro, Posse, on the valley of the rio Parahyba [rio Piabanha at Posse], 22 ° 23 ’ S, 43 ° 01 ’ W; C. Taylor, Thayer Expedition, 1865. MCZ 21113 (2, 114.8 – 253.0 mm SL): Mendez (Rio Paraíba do Sul or tributary at Mendes), 22 ° 32 ' S, 43 ° 44 ' W; C. Hartt & E. Copeland, 1865. Minas Gerais, rio Paraíba do Sul basin: MZUSP 42443 (2, 1 cs, 106.7 – 138.2 mm SL): Olaria, rio Pari, c. 21 ° 51 ’ S, 43 ° 52 ’ W; collector and date unknown. MZUSP 50805 (8, 128.8 – 274.5 mm SL); MZUSP 50803 (1, 58.9 mm SL): Torreões, rio do Peixe, near dam, 21 ° 53 ’ 15 ’’ S, 43 ° 32 ’ 29 ’’ W; F. A. Bockmann & A. Akama, 24 April – 1 May 1996. MZUSP 50800 (1, 80.5 mm SL): Torreões, rio do Peixe, near the bridge; F. A. Bockmann & A. Akama, 30 April 1996. MZUSP 50806 (2, 143.8 – 156.1 mm SL): Torreões, rio do Peixe at sítio da Jacutinga, “ fazenda do Tião Belo ”; F. A. Bockmann & A. Akama, 30 April – 1 May 1996. MZUSP 50804 (1, 64.7 mm SL): Minas Gerais, Torreões, córrego Jacutinga, tributary of rio do Peixe; F. A. Bockmann & A. Akama, 30 April 1996. MZUSP 89510 (1, 77.3 mm SL, cs): Torreões, rio do Peixe, c. 21 ° 53 ’ S, 43 ° 32 ’ W; F. A. Bockmann & A. Akama, Apr 1996. MZUSP 107915 (1, 227.0 mm SL): Juiz de Fora, rio do Peixe, UHE Picada, c. 21 ° 52 ' S, 43 ° 33 ’ W; C. Leal & M. Santos, 28 March 2009. MZUSP 83432 (1, 146.7 mm SL): Rio Preto, ribeirão Santana (trib. rio Paraíba do Sul), c. 22 ° 2 ’ S, 43 ° 46 ’ W; B. P. Nogueira & G. A. Pereira, 7 Dec 2002. MNRJ 14781 (1, 118.2 mm SL): Santana do Deserto, rio Cágado, trib. rio Paraibuna, 21 ° 59 ’ S, 43 ° 8 ’ W; D. F. Moraes Jr. et al., 29 Jun 1989. ZUEC 6380 (1, 139.7 mm SL): Rio Pomba, rio Pomba, 21 ° 18 ’ 0 ’’ S, 43 ° 12 ’ 18 ’’ W; D. M. Rosa, May 2010. Minas Gerais, rio Doce basin: MCZ 21130 (1, 241.1 mm SL): “ Rio Santo Antonio, Santa Anna de Ferros ” [Rio Santo Antonio (tributary of Rio Doce) at Ferros], 19 ° 13 ' S, 43 ° 1 ' W; T. Ward, 1865. MZUSP 100262 (2, 96.6 – 203.0 mm SL): Ferros, rio Santo Antônio, c. 19 ° 13 ' S 43 ° 1 ' W; C. Lima & H. Aguiar, 8 March 2008. MZUSP 51051 (1, 203.5 mm SL): Itambé do Mato Dentro, rio Preto do Itambé, 19 ° 24 ’ 40 ’’ S, 43 ° 18 ’ 41 ’’ W; J. C. Garavello et al., no date. MZUSP 59282 (3, 119.6 – 249.4 mm SL): Itambé do Mato Dentro, rio Preto do Itambé, trib. rio Santo Antônio, 19 ° 24 ’ 40 ’’ S, 43 ° 18 ’ 41 ’’ W; F. Vieira et al., 12 March 1994. MZUSP 59284 (3, 1 cs, 98.1 – 125.5 mm SL): Morro do Pilar, rio Preto, trib. rio Santo Antônio, 19 ° 14 ’ S, 43 ° 20 ’ W; F. Vieira et al., 13 March 1994. MZUSP 59285 (1, 93.9 mm SL): Itambé do Mato Dentro, rio Preto do Itambé, trib. rio Santo Antônio, 19 ° 24 ’ 40 ’’ S, 43 ° 18 ’ 41 ’’ W; F. Vieira et al., 27 Aug 1994. MZUSP 59281 (2, 89.1 – 199.3 mm SL); UFMG uncataloged (1, 120.5 mm SL): Morro do Pilar, rio Preto, trib. rio Santo Antônio, 19 ° 14 ’ S, 43 ° 20 ’ W; F. Vieira et al., 28 Aug 1994. MZUSP 59283 (1, 158.0 mm SL): Itambé do Mato Dentro, rio Preto do Itambé, trib. rio Santo Antônio, 19 ° 24 ’ 40 ’’ S, 43 ° 18 ’ 41 ’’ W; F. Vieira et al., 26 Nov 1994. MZUSP 59280 (2, 215.5 – 247.1 mm SL): Morro do Pilar, rio do Peixe, trib. rio Preto do Itambé; F. Vieira et al., 14 Sept 1996. MZUSP 59279 (3, 132.7 – 195.9 mm SL); UFMG uncataloged (1, 175.4 mm SL): São Sebastião do Rio Preto, rio Preto do Itambé, below rio do Peixe mouth; F. Vieira et al., 15 Sept 1996. MZUSP 59286 (5, 25.9 – 29.2 mm SL): rio Preto do Itambé, trib. rio Santo Antônio, 19 ° 24 ’ 40 ’’ S, 43 ° 18 ’ 41 ’’ W; F. Vieira et al., 25 Aug 1991. MZUSP 73104 (1, 232.6 mm SL): São Sebastião do Rio Preto, rio Santo Antônio, road to Córrego da Fonseca, approx. 11 km from Santo Antônio, c. 19 ° 16 ’ S, 43 ° 10 ’ W; F. Di Dário & S. Kakinami, 12 Sept 2001. MZUSP 73119 (1, 35.9 mm SL): Santo Antônio do Rio Abaixo, rio Santo Antônio, Benedito Martins beach, 19 ° 13 ’ 22 ’’ S, 43 ° 15 ’ 17 ’’ W; F. Di Dário, 14 Sept 2001. MZUSP 75309 (2, 55.3 – 68.3 mm SL); MZUSP 75310 (1, 82.5 mm SL): Conceição do Mato Dentro, rio Santo Antônio, fazenda das Campinas, road MG 0 10, km 158, 19 ° 4 ’ 56 ’’ S, 43 ° 25 ’ 53 ’’ W; F. Di Dário & B. Di Dário, 17 – 18 Dec 2001. UMMZ 216714 (1, 119.6 mm SL): “ Rio Doce: creeks, pools and river ” [= rio Xopotó at Alto rio Doce, 21 ° 1 ’ S, 43 ° 24 ’ W]; J. D. Haseman, 26 May 1908. MZUSP 59287 (1, 163.2 mm SL): Ponte Nova, rio Piranga, c. 20 ° 25 ’ S, 42 ° 55 ’ W; J. C. Garavello & A. S. Soares, 28 May – 4 June 1995. Rio Paraíba do Sul basin, no precise locality: UMMZ 216713 (4, 104.6 – 152.5 mm SL); UMMZ 216712 (4, 149.4 – 172.7 mm SL); MCZ 21119 (1, 77.3 mm SL); MCZ 21118 (3, 192.2 – 223.1 mm SL); MCZ 21120 and MCZ 21121 (10, 71.0 – 154.7 mm SL); NMW 62733 (3, 118.6 – 131.4 mm SL): “ Rio Parahyba ”; Thayer Expedition, 1865. Uncertain locality (see Remarks): MCZ 21104 (2, 59.2 – 62.2 mm SL): “ Rio Doce ” [Rio Doce, between Linhares and Aimorés], 19 ° 37 ' S 39 ° 49 ' W; C. F. Hartt & E. Copeland, 1865.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F3DFFA94EA4FBDBFEE2FA27.taxon	description	Chalceus opalinus (not Cuvier): Kner, 1860: 10 (“ Irisanga ”). Salmo pira-pitinga Kner (ex Natterer), 1860: 2 (name unavailable, published under the synonym of Chalceus opalinus).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F3DFFA94EA4FBDBFEE2FA27.taxon	diagnosis	Diagnosis. Brycon nattereri can be diagnosed from all remaining cis-andean Brycon species, with the exception of B. stolzmanni, B. coxeyi, B. coquenani, B. vermelha, B. insignis, B. howesi, B. dulcis, B. ferox, B. vonoi, and B. opalinus by possessing a color pattern consisting in a humeral blotch and a caudal peduncle blotch, without body stripes or other obvious color markings on caudal and anal-fins (vs. body stripes and caudal / anal fin color markings present; see Fig. 5). Brycon nattereri can be distinguished from B. ferox, B. vermelha, B. insignis, B. howesi, B. coquenani, and B. dulcis by possessing a rounded, obtuse head profile (vs. a distinctly acute head profile), and by possessing a caudal peduncle blotch extending into middle caudal-fin rays (vs. caudal peduncle blotch not extending into middle caudal-fin rays). Brycon nattereri can be distinguished from B. stolzmanni and B. coxeyi by the absence of a patch of dark pigmentation on the opercle (vs. dark patch of pigmentation present on opercle). Brycon nattereri can be additionally distinguished from B. insignis, B. howesi, B. coquenani, and B. vermelha by possessing a fifth infraorbital bone about as wide as high (vs. fifth infraorbital bone wider than high; see Fig. 6). Brycon nattereri can be distinguished from B. opalinus by possessing lateral line tubules with 2 – 5 branches (vs. tubules mostly simple), and caudal peduncle blotch extending into middle caudal-fin rays (vs. caudal peduncle blotch not extending into middle caudal-fin rays).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F3DFFA94EA4FBDBFEE2FA27.taxon	description	Description. Morphometric data are presented in Table 13. Middle-sized species, largest examined specimen 300.3 mm SL. Body slender to moderately slender. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly convex from latter point to basis of supraoccipital process, moderately to pronoucedly 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 moderately acute anteriorly, mouth terminal. Jaws isognathous to slightly anisognathous, premaxillary projecting slightly relative to dentary in some specimens, leaving outer row of premaxillary teeth exposed when mouth is closed. Maxillary moderately long, extending posteriorly to anterior margin to anterior third of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Five (1), 6 (7), 7 (6), 8 (18), 9 (16), 10 (16), 11 (4), 12 (3), or 14 (1) tricuspidate teeth in outer series. Three (4), 4 (37), 5 (23), or 6 (5) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 2 (20), 3 (48), or 4 (4) 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, both pentacuspidate. Maxillary with distal portion expanded and rounded in profile. Twelve to 26 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 8 (2), 9 (9), 10 (15), 11 (7), 12 (9), or 14 (2) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, tetra- to pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, tri- 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 16 – 24 small, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of fourth to sixth main series dentary teeth. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-six (2), 47 (4), 48 (2), 49 (11), 50 (11), 51 (15), 52 (12), 53 (6), 54 (5) ou 55 (6) scales in lateral line series. Laterosensory tube simple in specimens smaller than 95 mm SL, ramified in specimens larger than 100 mm SL. Tubules ramification increasing in complexity along ontogeny, specimens between 100 – 150 mm SL with tubules with two or three branches, larger specimens (> 180 mm SL) typically developing ascendant and descendant main branches and 2 – 5 secondary branches per scale. Horizontal scale rows between dorsal-fin origin and lateral line 8 (44) or 9 (28). Horizontal scale rows between lateral line and pelvic-fin 3 (2), 4 (49), or 5 (21). Circumpeduncular scales 15 (3), 16 (24), 17 (29), 18 (15), or 19 (3). Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 15 th (1) vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 18 (5), 19 (13), 20 (20), 21 (19), 22 (12), 23 (1), 24 (1), or 25 (1). First anal-fin pterygiophore inserting behind haemal spine of 26 th (1) vertebra. Last unbranched and anterior 4 – 5 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 20 per fin-ray main branch) small hooks on last unbranched and posterior main branch of branched rays 1 – 20, associated with dense, gelatinous tissue in a single specimen (CAS 79218, 204.2 mm SL). A single hook per ray segment, a few hooks at the anterior branch of some anal-fin rays. Sheath of scales covering basis of anal-fin rays composed of two scale rows, lower scale row formed by 16 – 21 rectangular scales. Pectoral-fin rays i, 12 (18), 13 (37), or 14 (18). Pelvicfin rays typically i, 7. One specimen (CAS 79218, 204.2 mm SL) with minute hooks on distal portion of posterior branch of branched pelvic-fin rays. Main caudal-fin rays 10 / 9. Caudal fin forked, lobes rounded. Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First branchial arch with 11 (1), 12 (11), 13 (5), or 14 (1) lower, 1 at angle, and 8 (1), 9 (7), 10 (7), or 11 (3) upper gill rakers. Vertebrae 46 (3). Supraneurals 10 (1). Coloration in alcohol. Top of head, snout, supraorbital, sixth infraorbital, and dorsal portion of body lightgrey to dark-brown. Second, third, fourth, and fifth infraorbitals, and opercle silvery except in specimens that lost most guanine, in which case brownish. Dentary, maxillary, gular area and lower portion of body light brown. Lateral portion of body silvery in specimens retaining guanine, brownish in specimens that were stored for a long period in formalin / alcohol. Humeral blotch present, relatively conspicuous in juveniles but little conspicuous in adult specimens, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of second, extending longitudinally to posterior margin of third to fourth lateral line scales, and vertically one and half scales high. Large, generally very conspicuous, oval-shaped caudal peduncle blotch, extending along 6 – 9 last lateral-line scales. Caudal peduncle blotch extending into middle 4 – 6 caudal fin-rays, often extending to the distal portion of middle caudal rays, forming a distinct caudal-fin stripe. Rayed fins clear, with dark chromatophores scattered over the interradial membranes. Adipose fin light-grey to light-brown. Juveniles (up to 100 mm SL) with narrow, wavy, irregularly-distributed vertical lines formed by dark pigmentation concentrated on anterior and posterior scales margins. Specimens from rio Cipó basin in upper rio São Francisco drainage (MZUSP 51052, MZUSP 58948, MZUSP 58949, MZUSP 58950) with overall body color, including fins, dark-brown, with scales on lateral portion of body with darkened margins (see item “ Variation ”, below). Color in life. Description based on pictures of several unpreserved specimens fished in several portions of the upper rio Paraná basin in Minas Gerais and upper rio Tocantins basin in Goiás, and from photographs of the specimens MZUSP 45793 and MNRJ 19893 immediatelly after collection. Dorsal area, top of head and snout lightgrey to light brown. Infraorbitals, opercular bones and sides of body silvery. Dark peduncular blotch very conspicuous. Fins (including adipose fin) yellowish to reddish. Common names. “ Peripetinga ” (Lütken, 1875, 2001: 222), “ pirapitinga ”, “ tabarana ”, “ tubarana ”. The name “ tabarana ” (from which “ tubarana ” is a variant) is more often employed for Salminus hilarii in southeastern Brazil.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F3DFFA94EA4FBDBFEE2FA27.taxon	distribution	Distribution. Distributed in headwater areas in the upper rio Paraná, upper rio Tocantins and upper rio São Francisco basins, central and eastern Brazil (Fig. 43). Bertoni (1939) listed the species as occurring in Paraguay. Though the occurrence of Brycon nattereri in the upper rio Paraná portion draining Paraguay is not unlikely, recent fish collections in that country (e. g. Géry & Mahnert, 1992) failed to obtain any specimens of this species, and this record is herein considered doubtful.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F3DFFA94EA4FBDBFEE2FA27.taxon	description	Variation. Specimens collected at the rio Cipó, a tributary of the rio das Velhas (rio São Francisco basin) present a general color pattern much more darker than the remaining specimens examined from the upper rio Paraná, upper rio Tocantins and other portions of the upper rio São Francisco basins. This overall dark pigmentation is very likely a phenotypical response to the dark waters of the rio Cipó (F. Vieira, pers. comm.), a fact also observed in the Brycon species occurring in the Amazon basin (see under B. amazonicus, below), and some other Brycon species from eastern Brazil (see species accounts of B. howesi, B. dulcis, and B. opalinus, above). Specimens from the upper rio Araguari, a tributary of the rio Paranaíba (upper rio Paraná basin) in Minas Gerais (MZUSP 44947 and MZUSP 45793) differ from remaining Brycon nattereri specimens by possessing a considerably convex predorsal profile, well-developed, wide infraorbitals 3 – 5, and a relatively ill-defined caudal peduncle blotch. There is not, however, a clear geographical pattern, since typical Brycon nattereri are known from the rio Araguari a little downstream of the area inhabited by these specimens. More material, as well as other type of data (e. g., molecular data) are necessary to understand what may lie behind this variation.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F3DFFA94EA4FBDBFEE2FA27.taxon	materials_examined	Ecological notes. Brycon nattereri is apparently restricted to clear- to dark-water, middle-sized rivers, with moderate to swift current, in headwater areas provided with well preserved riparian vegetation. Largest recorded size is 31 cm SL (Lima et al., 2008 b). A 51 cm SL and 2.6 kg specimen recorded as the maximum size for the species by Lima et al. (2008 a, b) from the Serra da Mesa area in the upper rio Tocantins was not examined by the author and more likely represents a misidentified specimen, perharps of the sympatric, large-sized Brycon gouldingi. Brycon nattereri has an omnivorous, highly opportunistic diet. In the upper rio Tocantins area, its diet was almost entirely composed of allochtonous items, especially seeds and fruits (though only fruits of Chrysobalanaceae and Caryocar sp. could be identified) and terrestrial arthropods (mainly beetles and ants). A lizard was recorded in the stomach of a 12 cm SL specimen. Smaller individuals tended to be more carnivorous, whereas larger specimens relied more heavily on plant material (Lima et al., 2008 b). Diet preferences of Brycon nattereri in the upper Paraná basin pointed to terrestrial and aquatic insects, detritus, small fish, and algae (Luz- Agostinho et al. 2006), in addition to fruits and other vegetal items (Lima et al., 2008 b). Smallest first maturation lengths recorded in the upper rio Tocantins basin were for females 11.4 and for males 9.8 cm SL, and in the upper rio Paraná basin 18.5 cm SL and 15.8 cm SL, respectively (Lima et al., 2008 b). Breeding individuals were detected during the middle dry season (May to July) in the upper rio Corumbá, rio Paraná basin and late dry season (June to August) in the upper rio Tocantins (Lima et al., 2008 a). In the upper rio Araguari basin (upper rio Paraná basin), however, breeding individuals were mainly detected in the late rainy season (March to April; Lima et al., 2008 b). A population of Brycon nattereri was reported to breed in a small river stretch, limited by waterfalls (Vieira et al. 2005), which strongly suggests that the species does not undertake long spawning migrations. Shortscale migratory behavior for the species was inferred from a seasonal fluctuation in abundance along the year in a tributary of the rio das Almas (upper rio Tocantins) (Vitorino Jr. et al., 2014). Very likely, the reproductive strategy of Brycon nattereri seems to fit the seasonal (periodic) type (sensu Winemiller & Rose, 1992). Juveniles (17 – 100 mm SL) are often collected in streams, which appear to be the recruitment areas for the species. Conservation. The decline of Brycon nattereri populations seems to be related to three factors, viz., deforestation of the riparian forest, water pollution, and dam building. Although deforestation and water pollution might be ascribed as the main causes of the decline of Brycon nattereri in areas historically more disturbed (e. g. the portion of the upper rio Paraná basin situated at the São Paulo state), construction of hydroelectric dams is a concurrent threat that has been increasing in the last decades. Many of the rivers of former known occurrence of the species are nowadays modified by hydroelectric dams and the species appears to subsist in those areas only at undisturbed tributaries, e. g., at the upper rio Tocantins basin in the Serra da Mesa dam area and at the upper rio Araguari basin in Nova Ponte dam area. Brycon nattereri is considered officially as threatened in Brazil, under the category “ vulnerable ” (see Lima et al., 2008 a, b, for additional information).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F3DFFA94EA4FBDBFEE2FA27.taxon	discussion	Remarks. Brycon nattereri was described by Günther (1864: 334), based on a specimen briefly described and identified by Kner (1860: 10) as Chalceus opalinus. This specimen (NMW 16396) was collected by J. Natterer at “ Irisanga ” (Rio Orissanga at Orissanga, now Estiva Gerbi, a tributary of rio Mogi-Guaçu, upper rio Paraná basin, São Paulo state; cf. Vanzolini, 1993). Kner (1860: 10) noticed that this specimen, in constrast with Cuvier’s (1819) description of Chalceus opalinus, possessed a conspicuous caudal peduncle blotch. This latter character was in fact the one used by Günther (1864), who has not examined Kner’s (1860) specimen, to diagnose it from Chalceus opalinus (“ coloration uniform, without blotchess ” versus “ a large black blotch at the base of the caudal fin ”; Günther, 1864: 334). Brycon nattereri possess, in fact, a caudal peduncle blotch considerably more conspicuous than the one found in B. opalinus, though there are some variation in this character (see item “ Variation ”, above). More importanly is the black pigmentation extending into the caudal-fin rays, a condition that clearly diagnoses Brycon nattereri from B. opalinus and, in fact, from all remaining, similar-looking Brycon species from eastern Brazil.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F3DFFA94EA4FBDBFEE2FA27.taxon	materials_examined	Material examined. Type material. NMW 16396 (1, 243.0 mm SL): " Irisanga " [= Brazil, São Paulo, Oriçanga (now Estiva Gerbi, 22 ° 16 ’ S, 46 ° 57 ’ W), rio Grande basin]; J. Natterer, no date (1822 – 1823) Holotype of Brycon nattereri Günther. MNHN 9589 (1, 130.2 mm SL); NMW 59761 (2, 115.9 – 192.0 mm SL); BMNH 1876.1.10.36 (1, 154.6 mm SL); USNM 44955 (1, 125.4 mm SL): “ Rio das Velhas ” [trib. Rio São Francisco, Minas Gerais, Brazil]. J. T. Reinhardt; syntypes of Brycon reinhardti Lütken. Non types. Brazil, upper rio Paraná basin, Goiás: MZUSP 58850 (1, 154.2 mm SL): Itumirim, rio Corrente, trib. rio Paranaíba, c. 18 ° 30 ’ 53 ’’ S, 52 ° 5 ’ 25 ’’ W; J. C. Garavello & M. L. Mussara, 28 Nov – 1 Dec 1994. MZUSP 43132 (1, 150.0 mm SL): Quirinópolis, rio Paranaíba system, c. 18 ° 34 ’ S, 50 ° 26 ’ W, collector and date not specifed. MNRJ 19895 (2, 172.7 – 196.5 mm SL): Catalão, córrego da Prata, trib. rio São Marcos, rio Paranaíba basin, 17 ° 47 ' 24 '' S, 47 ° 35 ' 46 '' W; F. A. Bockmann et al., 21 Nov 1999. MNRJ 19771 (1, 139.3 mm SL); MNRJ 19772 (1, 146.0 mm SL): Catalão, ribeirão São Domingos, trib. rio São Marcos, rio Paranaíba basin, 18 ° 5 ' 45 '' S, 47 ° 42 ' 10 '' W; F. A. Bockmann et al., 24 Sept 1999. MNRJ 19893 (1, 125.4 mm SL); MNRJ 19892 (5, 69.9 – 97.7 mm SL): Catalão, córrego da Anta Gorda, trib. rio São Marcos, fazenda Barreiro, 17 ° 52 ' 34 '' S, 47 ° 35 ' 46 '' W; F. A. Bockmann et al., 20 Nov 1999. MNRJ 19894 (1, 157.3 mm SL): Catalão, ribeirão Pires, trib. rio São Marcos, rio Paranaíba basin, 17 ° 52 ' 23 '' S, 47 ° 43 ' 45 '' W; F. A. Bockmann et al., 20 Nov 1999. MNRJ 19719 (1, 50.9 mm SL); MNRJ 19720 (1, 56.9 mm SL): Catalão, rio São Marcos, trib. rio Paranaíba, bridge at road GO- 506, Porto Carapina; C. A. Figueiredo et al., 22 Sept 1999. MNRJ 19710 (1, 39.0 mm SL): Catalão, córrego do Barreiro, trib. rio São Marcos, rio Paranaíba basin; C. A. Figueiredo et al., 24 Sept 1999. MNRJ 19891 (3, 22.0 – 23.2 mm SL): Catalão, ribeirão Buracão, trib. rio São Marcos, rio Paranaíba basin; F. A. Bockmann et al., 17 Nov 1999. ZUEC 6364 (1, 132.1 mm SL): Davinópolis, rio São Bento (trib. Rio São Marcos), 18 ° 8 ’ 7 ’’ S, 47 ° 38 ’ 24 ’’ W; T. C. Pessali, May 2010. MZUSP 51053 (3, 1 cs, 146.7 – 212.6 mm SL); MZUSP 68520 (1, 238.9 mm SL): Caldas Novas, rio Sapé, trib. rio Corumbá, rio Paranaíba basin, 17 ° 48 ’ S, 48 ° 35 ' W; NUPÉLIA, 23 – 24 March 1996. NUP 1304 (1, 242.7 mm SL): Caldas Novas, rio Corumbá, trib. rio Paranaíba, 17 ° 43 ' 37 '' S, 48 ° 32 ' 54 '' W; NUPÉLIA, 9 Apr 1999. MZUSP 104339 (2, 98.6 – 117.3 mm SL): Corumbaíba, córrego Libório, trib. rio Corumbá, 17 ° 55 ' 48 " S, 48 ° 28 ' 58 " W; NUPÉLIA, 21 March 1996. MZUSP 79967 (1, 126.6 mm SL): Luziânia, córrego Raso, trib. Rio Corumbá, 16 ° 19 ’ 28 ’’ S, 48 ° 11 ’ 43 ’’ W; H. L. R. Silva, no date. Distrito Federal: MNRJ 11265 (1, 114.7 mm SL): Brasília, rio Gama (rio São Bartolomeu basin), c. 15 ° 54 ’ S, 47 ° 54 ’ W; L. O. Soares, 1957. MZUSP 19861 (18, 19.3 – 26.4 mm SL); MZUSP 51007 (4, 17.6 – 25.6 mm SL): Planaltina, rio Piriripau (rio São Bartolomeu basin), c. 15 ° 41 ’ S, 47 ° 39 ’ W; Expedition MZUSP / USNM, 30 Sept 1977. IBGE uncat. (1, 300.3 mm SL): Brasília, Ribeirão Santa Maria, Lago de Santa Maria, Parque Nacional de Brasília, 15 ° 41 ’ 2 ’’ S, 48 ° 1 ’ 17 ’’ W; M. C. L. Ribeiro et al., 26 Aug 1994. IBGE uncat. (1, 111.8 mm SL): Brasília, córrego Três Barras, trib. Ribeirão Bananal, Parque Nacional de Brasília, 15 ° 39 ’ 48 ’’ S, 47 ° 57 ’ 21 ’’ W; M. C. L. Ribeiro et al., 19 Oct 1992. Minas Gerais: MZUSP 59002 (1, 140.1 mm SL): Uberlândia, rio Araguari, immediately downstream Miranda hydroelectric dam, 18 ° 54 ’ S, 48 ° 2 ’ W; V. Vono, Aug 1997. MZUSP 58910 (1, 206.9 mm SL): Nova Ponte, rio Araguari, Nova Ponte dam, c. 19 ° 7 ’ S, 47 ° 36 ’ W; F. Vieira et al., Feb 1994. MZUSP 59003 (1, 208.7 mm SL): Uberlândia, rio Araguari, UHE Miranda dam, c. 18 ° 54 ’ S, 48 ° 2 ’ W; V. Vono, May 1998. ZUEC 7940 (4, 98.6 – 106.7 mm SL): Veríssimo, rio Uberaba (trib. Rio Grande), 19 ° 41 ’ 46 ’’ S, 48 ° 11 ’ 30 ’’ W; D. C. Ribeiro & J. M. A. Chagas, 29 Sept 2012. MZUSP 44947 (2, 261.5 – 283.3 mm SL): Sacramento, rio Araguari, trib. rio Paranaíba, c. 19 ° 46 ’ S, 47 ° 20 ’ W; E. Zaniboni-Filho, 23 – 27 Sept 1991. MZUSP 45793 (2, 168.1 – 238.0 mm SL): Desemboque, rio das Velhas ou Araguari, 20 ° 0 ’ S, 47 ° 1 ’ W; A. L. Silva, July 1993. MZUSP 17834 (1, 52.9 mm SL): ribeirão da Prata, road São João Del Rei / Itutinga, c. 21 ° 13 ’ S, 44 ° 22 ’ W; H. A. Britski & J. C. Garavello, 10 Nov 1969. MZUSP 51526 (2, 199.8 – 216.7 mm SL): Baependi, rio Itaúna, Pedra Branca, c. 21 ° 57 ’ S, 44 ° 51 ’ W; M. A. F. Filho, 9 Feb 1996. LBP 8101 (1, 236.0 mm SL): Carrancas, rio Capivari, c. 21 ° 32 ’ S, 44 ° 42 ’ W; C. Oliveira et al., 18 May 2009. São Paulo: MZUSP 3359 (1, 99.6 mm SL): Monte Alegre do Sul, rio Camanducaia, c. 22 ° 42 ’ S, 46 ° 40 ’ W; J. Lima, 1943. CAS 79218 (1, 204.2 mm CP), Itatiba [Rio Atibaia at Itatiba, c. 22 ° 58 ’ S, 46 ° 50 ’ W]; J. Lima, 1907. MNRJ 20159 (4, 68.3 – 92.1 mm SL): Itararé, rio Verde road SP- 258, Itararé / Itapeva border, 24 ° 5 ’ 2 ’’ S, 49 ° 12 ’ 2 ’’ W; P. A. Buckup & M. R. S. Melo, 25 Jan 2000. Paraná: MZUEL 826 (2, 222.9 – 253.0 mm SL): Tibagí, Guartelá Ecological State Reserve, rio Iapó (trib. rio Tibagí), c. 24 ° 32 ’ S, 50 ° 17 ’ W; O. A. Shibatta et al., 1 Nov 1996. MZUEL 5746 (2, 190.0 – 230.0 mm SL): Santa Amélia, ribeirão Laranjinha (trib. rio das Cinzas), 23 ° 17 ’ 52 ’’ S, 50 ° 28 ’ 43 ’’ W; B. Galindo et al., 27 Feb 2011. NUP 8534 (1, 151.0 mm SL): Turvo, rio Pedrinho, trib. rio Ivaí, 25 ° 4 ’ S, 51 ° 30 ’ W; D. Viana, 20 Jan 2009. Rio São Francisco basin, Minas Gerais: MZUSP 58949 (2, 191.4 – 238.4 mm SL): Jaboticatubas / Santana do Riacho, rio Cipó (trib. Rio das Velhas), Serra do Cipó National Park, c. 19 ° 20 ’ S, 43 ° 38 ’ W; F. Vieira et al., 31 May 1994. MZUSP 58948 (1, 111.7 mm SL): Jaboticatubas / Santana do Riacho, rio Cipó (trib. Rio das Velhas), Serra do Cipó National Park, c. 19 ° 20 ’ S, 43 ° 38 ’ W; F. Vieira et al., 15 March 1994. MZUSP 58950 (1, 244.7 mm SL): Jaboticatubas / Santana do Riacho, rio Cipó (trib. Rio das Velhas), Serra do Cipó National Park, c. 19 ° 20 ’ S, 43 ° 38 ’ W; F. Vieira et al., 30 Nov 1994. MZUSP 51052 (1, 228.4 mm SL): Jaboticatubas / Santana do Riacho, rio Cipó (trib. Rio das Velhas), Serra do Cipó National Park, c. 19 ° 20 ’ S, 43 ° 38 ’ W; F. Vieira et al., Aug 1991. MZUSP 1599 (1, 217.0 mm SL): Lassance, ribeirão São José (trib. Rio das Velhas), Serra do Cabral, c. 17 ° 54 ’ S, 44 ° 31 ’ W; E. Garbe, 1912. MNRJ 11264 (1, 64.0 mm SL): Pirapora, córrego da Cachoeira, c. 17 ° 20 ’ S, 44 ° 59 ’ W; G. S. Myers, P. M. Ribeiro & A. L. Carvalho, Oct 1942. ZUEC 6383 (2, 51.8 – 63.5 mm SL): Buritizeiro, ribeirão das Gaitas, trib. rio do Sono, 17 ° 17 ’ 43 ’’ S, 45 ° 18 ’ 50 ’’ W; T. C. Pessali & T. A. Barroso, Nov 2011. MZUSP 17019 (1, 122.5 mm SL): Buritis, ribeirão dos Confins (trib. Rio Urucuia), 15 ° 38 ’ S, 46 ° 22 ’ W; P. E. Vanzolini, Oct 1964. UFRGS 11266 (5, 54.6 – 67.7 mm SL): Unaí / Palmeirinha, trib. rio Salobro at road Unaí / Palmeirinha, 16 ° 09 ’ 24 ’’ S 46 ° 44 ’ 48 ’’ W, G. Frainer, F. R. Carvalho & V. A. Bertaco, 11 Sept 2009. Rio Tocantins basin, Goiás: MZUSP 113836 (1, 75.3 mm SL): Goiás, São João da Aliança, ribeirão das Brancas, road GO- 118, 14 ° 36 ' 56.89 " S 47 ° 29 ' 55.94 " W; F. Dagosta et al., 27 Nov 2013. MZUSP 113865 (7, 94.4 – 115.5 mm SL): Teresina de Goiás, rio das Pedras, Cachoeira do Poço Encantado, 13 ° 52 ' 28.90 " S, 47 ° 15 ' 45.20 " W; F. Dagosta et al., 29 Nov 2013. MZUSP 113885 (3, 79.4 – 146.5 mm SL): São Jorge, rio São Miguel, at meeting with rio Segredo, 14 ° 8 ' 15.00 " S 47 ° 39 ' 58.00 " W; F. Dagosta et al., 28 Nov 2013. MZUEL 5745 (4, 19.6 – 32.0 mm SL): Pirenópolis, ribeirão Santa Maria (trib. rio das Almas), 15 ° 50 ’ 50 ’’ S, 48 ° 55 ’ 8 ’’ W; O. A. Shibatta & A. Claro, 15 Jul 2010. MNRJ 13069 (1, 89.2 mm SL): Cavalcante, córrego São Bento, trib. rio Preto (trib. rio Tocantinzinho); D. F. Moraes Jr., 9 Dec 1987. MCP 23938 (6, 22.2 — 34.0 mm SL): Cavalcante, córrego Móquem, road Colinas / Cavalcante, 13 ° 56 ' 28 ' ’ S, 47 ° 42 ' 31 ' ’ W; W. R. Koch et al., 5 Aug 1996. CAS 68822 (2, 197.3 – 235.6 mm SL): Córrego do Poichos into R. Maranhão; C. Ternetz, 10 Nov 1923. CAS 68828 (1, 113.2 mm SL); CAS 68828 (2, 172.1 – 232.2 mm SL): córrego do Monjolo into rio Maranhão into rio Tocantins; C. Ternetz, 1 – 14 Oct 1923. CAS 68831 (3, 126.8 – 214.0 mm SL): Córrego dos Bichos into R. Mangabeira into R. Maranhão; C. Ternetz, 14 Nov 1923. MZUSP 95051 (1, 83.4 mm SL): Minaçu, trib. Rio Tocantizinho, near Serra da Mesa dam, 14 ° 8 ’ 46 ’’ S, 48 ° 2 ’ 52 ’’ W; O. T. Oyakawa & K. C. Mautari, March 2006. MZUSP 62268 (2, 57.7 – 68.9 mm SL): Minaçú, trib. rio Tocantizinho, 14 ° 04 ’ 43 ’’ S, 48 ° 12 ’ 49 ’’ W; M. T. Rodrigues et al., 19 Dec 1995. MZUSP 59623 (3, 1 cs, 84.1 – 144.6 mm SL): Minaçú, trib. rio Tocantizinho, 14 ° 02 ' 27 '' S, 48 ° 12 ' 22 '' W; M. T. Rodrigues et al., 20 Dec 1995. MZUSP 104124 (1, 212.0 mm SL); MZUSP 104223 (1, 179.4 mm SL): Minaçú, rio Tocantizinho, 14 ° 12 ’ 44 ’’ S, 48 ° 3 ’ 24 ’’ W; D. F. Moraes & D. A. Halboth, 9 – 16 Dec 1995. MZUSP 104125 (4, 98.4 – 220.2 mm SL): Minaçú, rio Tocantizinho, 14 ° 12 ’ 44 ’’ S, 48 ° 3 ’ 24 ’’ W; D. F. Moraes & D. A. Halboth, 17 Apr 1996. MZUSP 103126 (4, 125.7 – 206.4 mm SL): Minaçú, rio Tocantizinho, 14 ° 12 ’ 44 ’’ S, 48 ° 3 ’ 24 ’’ W; D. F. Moraes & D. A. Halboth, 11 Aug 1996. MZUSP 104222 (1, 204.3 mm SL): rio Tocantizinho, mouth of rio Palmeirinha, 14 ° 3 ’ 57 ’’ S, 48 ° 29 ’ 37 ’’ W; D. F. Moraes & D. A. Halboth, 16 Feb 1995. Distrito Federal: IBGE uncat. (4, 94.0 – 187.0 mm SL): Brasília, Ribeirão Palmeiras, trib. rio Maranhão, 15 ° 31 ’ 10 ’’ S, 47 ° 43 ’ 40 ’’ W; M. C. L. Ribeiro et al., 8 Jul 1998. IBGE uncat. (2, 138.0 – 165.0 mm SL): Brasília, Córrego Monjolo, trib. rio da Palma (trib. rio Maranhão), 15 ° 31 ’ 9 ’’ S, 47 ° 58 ’ 9 ’’ W; M. C. L. Ribeiro et al., 23 Jul 1998. IBGE uncat. (10, 85.0 – 173.0 mm SL): Brasília, Ribeirão Dois Irmãos, trib. rio do Sal (trib. rio Maranhão), 15 ° 31 ’ 9 ’’ S, 48 ° 6 ’ 56 ’’ W; M. C. L. Ribeiro et al., 12 Aug 1998.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F26FFA14EA4F9CEFBBAFEFA.taxon	description	Chalceus carpophagus (not Valenciennes): Castelnau, 1855: 68, pl. 34, fig. 3 (description; “ Rio de Sabara de la province de Minas Geraës ”).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F26FFA14EA4F9CEFBBAFEFA.taxon	diagnosis	Diagnosis. Brycon orthotaenia can be distinguished from all remaining cis-andean Brycon species, except for B. orbignyanus, B. hilarii, B. whitei, and B. polylepis, by possessing a caudal peduncle blotch extending as a stripe to the distal portion of caudal-fin rays (caudal peduncle blotch, when present, limited to the caudal peducle or extending only into centralmost caudal-fin rays; Fig. 5). It can be distinguished from Brycon hilarii and B. whitei primarily by possessing lower scale counts (49 – 58, modally 52 lateral line scales, vs. 67 – 82, modally 74 in B. hilarii and 66 – 76, modally 70 in B. whitei; 9 – 12, modally 10 scales between lateral line and dorsal-fin basis, vs. 12 – 17, modally 15 in B. hilarii and 12 – 13, modally 13, in B. whitei; 18 – 21, modally 19 circumpeduncular scales, vs. 20 – 28, modally 26, in B. hilarii, and 19 – 24, modally 21, in B. whitei). Brycon orthotaenia can be further distinguished from B. whitei by lacking a midlateral dark stripe (vs. midlateral dark stripe present). Brycon orthotaenia can be distinguished from B. orbignyanus by possessing a aproximately rounded, obtuse head profile (versus pointed in the latter species; compare Fig. 6 F and H), by possessing a relatively broad and short dentigerous premaxilary surface (vs. a narrow and elongated dentigerous surface of the premaxillary), by possessing a lower number of teeth in the second, inner premaxilary row (not counting the teeth of the second row situated between the first and third rows) (3 – 5, modally 5, versus 5 – 9, modally 6, in B. orbignyanus), and by possessing the anteriormost four dentary teeth considerably larger than the remaining teeth (vs. dentary teeth decreasing gradualy in size in B. orbignyanus).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F26FFA14EA4F9CEFBBAFEFA.taxon	description	Description. Morphometric data are presented in Table 14. Middle-sized species, largest examined specimen 392.8 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, slightly concave from latter point to basis of supraoccipital process, moderately to pronoucedly 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 to moderately 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 obtuse, mouth terminal. Jaws isognathous to slightly anisognathous, premaxillary projecting slightly relative to dentary, leaving outer row of premaxillary teeth and in some specimens a portion of second series exposed when mouth is closed. Maxillary moderately long, extending posteriorly to anterior margin to anterior third of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Eight (6), 9 (23), 10 (15), 11 (11), 12 (5), or 13 (1) tricuspidate, relatively small teeth in outer series. Three (2), 4 (26), 5 (31), or 6 (1) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 2 (20), 3 (49), or 4 (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, penta- to hexacuspidate. Maxillary margins approximately parallel, straight in profile. Ten to 22 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 6 (2), 7 (4), 8 (21), 9 (19), 10 (6), 11 (3), or 12 (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 – 17 small, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of sixth to seventh main series dentary teeth. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-nine (3), 50 (2), 51 (7), 52 (11), 53 (13), 54 (8), 55 (8), 56 (9), 57 (5), or 58 (1) 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 between 100 – 150 mm SL with tubules with two or three branches, four to seven branches in specimens between 150 – 250 mm SL, and with more than 10 branches and developing a dendritic pattern of ramification, with tubules overlapping each other in larger (> 300 mm SL) specimens. Horizontal scale rows between dorsal-fin origin and lateral line 9 (1), 10 (33), 11 (30), or 12 (2). Horizontal scale rows between lateral line and pelvic-fin 5 (28), 6 (37), or 7 (1). Circumpeduncular scales 18 (10), 19 (44), 20 (9), or 21 (1). Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 13 th (1), 14 th (1), or 15 th (1) vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 22 (1), 23 (1), 24 (8), 25 (16), 26 (21), 27 (16), 28 (4), or 29 (1). First anal-fin pterygiophore inserting behind haemal spine of 25 th (1) or 26 th (1) vertebra. Anal-fin rays decreasing only slightly in size towards anal-fin end. Anal fin displaying several (c. 8 – 15 per fin-ray main branch) small hooks on last unbranched and posterior main branch of branched rays 1 – 12, associated with dense, gelatinous tissue in 8 specimens (MZUSP 39278, 6, 220.2 – 247.7 mm SL; MZUSP 17018, 1, 206.8 mm SL; MZUSP 2007, 1, 193.1 mm SL). A single hook per ray segment. Sheath of scales covering basis of anal-fin rays composed of four scale rows, lower scale row formed by 25 – 28 rectangular scales. Pectoral-fin rays i, 11 (2), 12 (25), 13 (40), or 14 (2). Pelvicfin rays i, 7. Main caudal-fin rays 10 / 9. Caudal fin slightly forked, distal margin slightly concave. Central caudalfin rays 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 at the middle caudal-fin projection. 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 14 (2), 15 (1), 16 (12), 17 (14), 18 (6), or 19 (1) lower, 1 at angle, and 15 (1), 16 (6), 17 (14), 18 (12), or 19 (4) upper gill rakers. Vertebrae 46 (4). Supraneurals 10 (4). Coloration in alcohol. Top of head, snout, supraorbital, and sixth infraorbital gray to light-brown. Dorsal portion of body light-brown to dark-brown, with silvery hue in specimens retaining guanine. Second, third, fourth, and fifth infraorbitals, and opercle silvery. Dentary, maxillary, gular area and lower portion of body clear in relatively recently collected specimens, light brown in specimens stored for some years in alcohol. Lateral portion of body silvery in relatively recently collected specimens, light brown, with a silvery hue, in specimens that were stored for a long period in formalin / alcohol. Humeral blotch present, conspicuous, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of second, extending longitudinally to posterior margin of fourth to fifth, 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. Caudal peduncle with broad median stripe, originating 4 – 6 scales from hypural joint and continuing posteriorly over 4 central principal caudal-fin rays to caudal-fin distal margin. Remaining caudal-fin rays, and remaining fins, clear. Color in life. Description based on the examination of several living specimens collected at the middle rio São Francisco at São Romão, one of which preserved (MZUSP 94993), and on a picture of one unpreserved specimen, fished at the Três Marias dam. Lateral body surfaces, infraorbital and opercular bones silvery. Dorsum and adiposefin light grey, with a greenish tinge. Anal, pelvic, and pectoral-fins pinkish. Caudal fin pinkish to deep red, basis of caudal fin yellowish. Sexual dimorphism. Eight examined specimens (MZUSP 39278, 6, 220.2 – 247.7 mm SL; MZUSP 17018, 1, 206.8 mm SL; MZUSP 2007, 1, 193.1 mm SL) presenting numerous small hooks present on first unbranched and posterior branch of branched anal-fin rays. Two of these specimens were dissected and proved to be males. Female specimens identified through dissection or through presence of ripe eggs at the urogenital opening (MZUSP 39728, 2, 190.3 – 245.8 mm SL) lack fin hooks. Common names. “ Matrinchã ” (Günther, 1864: 335, como “ matrinxim ”; Lütken, 1875: 223; 2001: 138); “ piabanha ” (Castelnau, 1855: 68, as “ piabana ”). Quoting Reinhardt, in Lütken, 2001: 138, footnote 1, “ this species is incorrectly called “ Piabanha ”, a name which actually belongs to another fish species that does not occur at the Rio das Velhas ” (our translation). Although “ piabanha ” is a common name widely employed for several Brycon species, the common name currently in use by fishermen for B. orthotaenia in the rio São Francisco basin is generally “ matrinchã ”.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F26FFA14EA4F9CEFBBAFEFA.taxon	distribution	Distribution. Endemic from the rio São Francisco basin, eastern Brazil (Fig. 52). There is a purported record for the species from the rio Poti, a tributary of the rio Paranaíba basin in northeastern Brazil (MCZ 21268). These two specimens were collected by Orestes St. John, a member of the Thayer Expedition, in December 1865. There are no other records for the species for the rio Parnaíba and in fact, so far no Brycon species has been reported from this river basin (see the item Biogeography, below). Since Orestes St. John also collected at the rio São Francisco basin (Higuchi, 1996), we consider that this specimens were probably mislabelled and that they were actually collected at the rio São Francisco basin.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F26FFA14EA4F9CEFBBAFEFA.taxon	discussion	Remarks. Günther (1864: 335) described Brycon orthotaenia based on a single specimen, collected at the “ River Cipo ” in Brazil. The description is quite short, with no figures appended. Lütken (1874: 135 – 136) described Brycon lundii from the rio das Velhas. In this description (attributed to Reinhardt), he notices that B. lundii might be actually a synonym of B. orthotaenia, both nominal species differing only in lateral-line scale counts (slightly higher in B. lundii) and in the absence of the inner dentary pair of symphyseal teeth in the holotype of B. orthotaenia. A little later, Lütken (1875: 225; 2001: 114) discussed in more detail the possible synonym between both nominal species, noticing that the distinct scale counts were probably a result of different count methods employed by him and Günther, and that the inner dentary teeth might have been simply lost in the holotype of Brycon orthotaenia. However, these remarks were overlooked by subsequent authors working with specimens from the rio São Francisco basin, which generally prefered to use Lütken’s name (Steindachner, 1917; Amaral Campos, 1950; Britski et al., 1984). Brycon orthotaenia was relegated as a synonym of B. orbignyanus by Berg (1895: 124), based on the erroneous assumption that the type locality of B. orthotaenia was at the La Plata basin. This synonym was reproduced in latter compilations on the genus (e. g., Fowler, 1950). Howes (1982: 33, 38 – 40) reexamined the holotype of Brycon orthotaenia, plus additional material of both B. orthotaenia and B. orbignyanus, and argued that B. orthotaenia was not a synonym of B. orbignyanus, but instead a valid species and very probably a senior synonym of B. lundii, a point of view also supported by Géry & Mahnert (1992: 817). Howes (1982: 39) incorrectly remarked that the “ River Cipo ” is a tributary of the rio Itapicurú (an independent coastal river system in northern Bahia state, northeastern Brazil). However, the rio Cipó is actually a well-known tributary of the rio das Velhas (a tributary of the rio São Francisco) at Minas Gerais state, a region in fact visited by the collector of the holotype of Brycon orthotaenia, Cumberland (Lütken, 1875: 134, 225; 2001: 29, 114), and, oddly, the very same river basin from where the type-material of B. lundii was collected. Although neither the holotype of Brycon orthotaenia (BMNH 1861.5.16.71) nor the syntypes of Brycon lundii (ZMUC 227, ZMUC 228 - 229, ZMUC 232; though Lütken, 1875: 225; 2001: 112, mentioned that he had a single specimen of this species) were directly examined in the present study, there is no doubt that, as earlier advanced by Lima (2003: 177 – 178) that both nominal species are synonymous. The examination of a photograph of the holotype of Brycon orthotaenia, the good description of B. lundii provided by Lütken (1875, 2001), and a wealthy of specimens from the entire rio São Francisco examined in the present study allow us to conclude that the differences between Günther’s and Lütken’s description are in fact ascribable to different lateral-line scale count methods and loss of the inner symphyseal dentary teeth in the holotype of B. orthotaenia. Valenciennes (1850: 252 – 253) described Chalceus carpophaga based on three syntypes, one of which said to be collected at “ l’Amazone ” by Castelnau. Castelnau (1855: 68), when referring to this specimen, mentioned that it was actually collected at the “ rio de Sabara de la province de Minas Geraës ”. The “ rio de Sabara ” is the rio das Velhas, thus this specimen was actually collected at the same river basin where the type specimens of Brycon orthotaenia and B. lundii were collected. Lütken (1875: 225 – 226; 2001: 114 – 115) considered Castelnau’s (1855) Chalceus carpophagus as probably distinct from Brycon orthotaenia and B. lundii. Howes (1982: 17) considered that Castelnau’s syntype of Chalceus carpophagus was not conspecific with the remaining two syntypes but instead was probably a specimen of B. orthotaenia. Géry & Mahnert (1992: 816) selected as lectotype of Chalceus carpophagus the syntype from the Essequibo River, and consequently the syntype collected by Castelnau at the rio das Velhas (MNHN A. 8615) became a paralectotype. Géry & Mahnert (1992: 816 – 817) compared the data of this specimen with the data of the type material of both Brycon orthotaenia and B. lundii and agreed with Howes (1982) in considering it as probably a specimen of B. orthotaenia. The examination of this paralectotype allowed us to confirm that it is in fact a specimen of Brycon orthotaenia. Ecological notes. At the rio São Francisco basin, Brycon orthotaenia is sometimes found syntopically with the its congener B. nattereri, but contrary to this species it favors larger, low-gradient rivers. It is also commonly found in floodplain lakes (Pompeu & Godinho, 2003 a; Pompeu & Godinho, 2006). As other Brycon species, B. orthotaenia is recorded to be an omnivore, with a tendency towards herbivory. Lütken (1875: 224; 2001: 114) reported that Reinhardt found vegetal matter and seeds in two stomach contents. Pompeu & Godinho (2003 b) examined 30 stomachs of Brycon orthotaenia specimens collected in marginal lagoons of the rio São Francisco and found it to be composed mainly of macrophytes and filamentous algae, with smaller amounts of aquatic insects, seeds and fruits. As most of its congeners, Brycon orthotaenia is a highly-fecund, total-spawner migratory fish (Sato et al., 2003), with a mean total fecundity of 400.000 eggs (Sato, Fenerich-Verani & Godinho, 2003). The breeding season is recorded in the rio São Francisco at Pirapora to be between October and January, with smallest recorded sizes of first maturity reported as being 32.0 cm TL for males and 40.5 cm TL for females (Gonçalves et al., 2003). Females are reported to grow larger than males and a maximum weigth of 7 kg is recorded for the species (Sato & Godinho, 2003). An analysis of microsatellites loci in populations of B. orthotaenia from below Três Marias dam by Sanches et al. (2012) indicated the presence of two distinct populations, which might indicate distinct sites of communal reproduction (“ piracema ”) for migratory schools of each population within a relatively small stretch of the rio São Francisco basin.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F26FFA14EA4F9CEFBBAFEFA.taxon	description	Conservation. Brycon orthotaenia is an important target of subsistence and small-scale commercial fisheries in the rio São Francisco basin (Sato & Godinho, 2003). It was the third more captured species, and the fifth most important fish species in biomass in the fisheries of the rapids of Pirapora during 1999 (Godinho et al., 2003). Though reported to be declining in the last decades (e. g., Sato & Godinho, 2003), it is still a common and widespread species in the undammed portion of the rio São Francisco river, i. e., between the Três Marias and Sobradinho reservoirs.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F26FFA14EA4F9CEFBBAFEFA.taxon	materials_examined	Material examined. Type material. BMNH 1861.6. 16.71, 330 mm SL: “ River Cipo ” [= Rio Cipó, trib. Rio das Velhas, Minas Gerais, Brazil, c. 18 ° 41 ’ S, 43 ° 59 ’ W]; Cumberland, no date (photographs only). Holotype of Brycon orthotaenia Günther. MNHN A. 8615 (1, 314.1 mm SL): " Rio de Sabara de la province de Minas Geraës " (= Brazil, Minas Gerais, rio das Velhas at Sabará, 19 ° 53 ' S, 43 ° 48 ' W); F. Castelnau. Paralectotype of Chalceus carpophagus Valenciennes (designated by Géry & Mahnert, 1992: 816). Non types. Brazil, rio São Francisco basin. Minas Gerais: MZUSP 18950 (5, 173.8 – 248.0 mm SL): Três Marias reservoir, rio São Francisco, c. 18 ° 22 ’ S, 45 ° 17 ’ W; CODEVASF, 1978. MZUSP 95402 (35, 3 sc, 90.5 – 230.8 mm SL): Três Marias, CODEVASF fish hatchery; O. T. Oyakawa et al., 4 Oct 2007. MZUSP 95166 (2, 153.0 – 153.8 mm SL): Três Marias, rio São Francisco, immediately downstream Três Marias reservoir, c. 18 ° 10 ’ S, 45 ° 14 ’ W; Y. Sato, 23 March 2007. MCP 14119 (5, 175.8 – 219.4 mm SL): Três Marias, rio São Francisco, between Três Marias and Pirapora; Y. Sato, Nov 1987. MZUSP 1619 (1, 102.2 mm CP): Pirapora, rio São Francisco, 17 ° 21 ’ S, 44 ° 57 ’ W; E. Garbe, 1903. MNRJ 47676 (1, 392.8 mm SL): Pirapora, rio São Francisco, 17 ° 21 ’ S, 44 ° 57 ’ W; G. S. Myers, P. Miranda-Ribeiro & A. L. Carvalho, Oct 1942. MZUSP 73836 (1, 275.0 mm SL): Lassance, rio das Velhas at the ferry, 17 ° 54 ’ 45 ’’ S, 44 ° 34 ’ 20 ’’ W; C. B. M. Alves & P. S. Pompeu, 17 Jun 1999. MCZ 21268 (1, 167.2 mm SL): Rio das Velhas (precise locality unknown); O. St. John, 1865. MZUSP 89514 (1 skel., 181.6 mm SL): Pirapora, rio São Francisco, between Buritizeiro and Pirapora; A. Akama et al., 28 Aug 2004. MZUSP 39728 (19, 181.2 – 319.3 mm SL): rio São Francisco and tributaries, immediately below Pirapora, c. 17 ° 4 ’ S, 44 ° 57 ’ W; Y. Sato et al., Nov 1987 – Aug 1988. MZUSP 39693 (1, 230.7 mm SL), rio São Francisco, Pontal do Abaeté, c. 17 ° 4 ’ S, 44 ° 57 ’ W; Y. Sato et al., 22 Jul 1988. MZUSP 94993 (1, 141.2 mm SL): São Romão, rio São Francisco, near Ribanceira village, 16 ° 28 ’ 26 ’’ S, 45 ° 6 ’ 16 ’’ W; F. C. T. Lima & M. Ribeiro, 23 – 25 Jun 2007. MNRJ 18130 (1, 323.0 mm SL): Palmital, rio Preto, trib. rio Paracatu, below Cachoeira de Queimados, Fazenda Mata, 16 ° 12 ’ 35 ’’ S, 47 ° 13 ’ 54 ’’ W; C. A. Figueiredo & D. F. Moraes Jr., 8 Jan 1998. MZUSP 17018 (1, 206.8 mm SL): Buritis, ribeirão Confins (trib. rio Urucuia), 15 ° 38 ’ S, 46 ° 22 ’ W; P. E. Vanzolini, Oct 1964. ANSP 171719 (1, 162.9 mm SL): Rio Empoeirado (lagoon) on road bewtween Pedra de Maria and Januária, 15 ° 33 ’ 56 ’’ S, 44 ° 24 ’ 6 ’’ W; S. A. Schaefer et al., 13 Jul 1993. MNRJ 15834 (2, 188.2 – 189.7 mm SL): Itacarambi, rio São Francisco, c. 15 ° 6 ’ S, 44 ° 5 ’ W; D. F. Moraes Jr. & J. C. Oliveira, 17 Aug 1990. MZUSP 42074 (1, 120.7 mm SL); MNRJ 15866 (1, 122.2 mm SL): Manga, lagoon at Mocambinho, rio São Francisco, c. 15 ° 4 ’ S, 44 ° 1 ’ W; J. C. Oliveira & O. T. Oyakawa, 17 – 22 Jul 1990. MNRJ 16065 (1, 139.9 mm SL): Manga, lagoa do Sossego, Mocambinho, c. 15 ° 4 ’ S, 44 ° 1 ’ W; D. F. Moraes Jr. & J. C. Oliveira, 23 Aug 1990. MNRJ 14226 (4, 104.0 – 116.6 mm SL): Manga, rio Mocambinho, at mouth at the rio São Francisco, c. 15 ° 4 ’ S, 44 ° 1 ’ W; D. F. Moraes Jr. & J. C. Oliveira, 17 Aug 1990. MNRJ 16345 (2, 100.6 – 105.1 mm SL): Manga, riacho Mocambinho, trib. rio São Francisco, Mocambinho, c. 15 ° 4 ’ S, 44 ° 1 ’ W; J. C. Oliveira et al., 6 Apr 1990. MNRJ 16352 (1, 111.7 mm SL): Manga, lagoa de Mocambinho, rio São Francisco right margin, c. 15 ° 4 ’ S, 44 ° 1 ’ W; J. C. Oliveira, 6 March 1990. MNRJ 15856 (1, 138.5 mm SL), Manga, rio São Francisco at ilha do Caju, below Mocambinho, c. 15 ° 2 ’ S, 44 ° 0 ’ W; D. F. Moraes Jr. & J. C. Oliveira, 21 Aug 1990. MNRJ 15870 (1, 156.1 mm SL): Manga, lagoa do Cajueiro (rio São Francisco, right margin), below Mocambinho, c. 15 ° 2 ’ S, 44 ° 0 ’ W; D. F. Moraes Jr. & L. C. Alvarenga, Sept 1990. MNRJ 16191 (1, 160.6 mm SL): Manga, lagoa do Caju, rio São Francisco right margin, 4 km below Mocambinho, c. 15 ° 2 ’ S, 44 ° 0 ’ W; D. F. Moraes & J. C. Oliveira, 23 Aug 1990. LISDEBE uncat. (1, 210.0 mm SL): Manga, rio Verde, trib. rio São Francisco, 5 km from Gado Bravo, c. 14 ° 44 ’ S, 43 ° 49 ’ W; G. B. Santos, 7 April 1986. MNRJ 16193 (2, 127.7 – 134.1 mm SL): rio Verde Grande, 5 km from its mouth at the rio São Francisco, Minas Gerais / Bahia border, c. 14 ° 36 ’ S, 43 ° 52 ’ W; D. F. Moraes & J. C. Oliveira, 23 Aug 1990.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F26FFA14EA4F9CEFBBAFEFA.taxon	description	Bahia: ZUEC 9189 (1, 168.0 mm SL): Carinhanha, rio Carinhanha (marginal lagoon), 14 ° 20 ’ 46 ’’ S, 43 ° 47 ’ 26 ’’ W; G. N. Salvador & E. Estevam, 6 Oct 2014. MCP 16676 (1, 157.1 mm SL): Bom Jesus da Lapa, riacho Santana, 31 km S from Bom Jesus da Lapa, road to Malhada, 13 ° 31 ’ 13 ’’ S, 43 ° 21 ’ 28 ’’ W; R. E. Reis et al., 18 Jul 1993. UFPB 2956 (3 of 8, 1 cs, 89.0 – 108.4 mm SL): riacho Pedra Branca, trib. rio Corrente, 31 km E from Santa Maria da Vitória, c. 13 ° 18 ’ S, 43 ° 51 ’ W; G. G. Filho & R. S. Rosa, 6 April 1994. MZUSP 94656 (1, 151.5 mm SL): Santa Maria da Vitória, rio Corrente at Santa Maria da Vitória, 13 ° 24 ’ S, 44 ° 11 ’ W; O. T. Oyakawa et al., 6 May 2007. MZUSP 28796 (1, 154.1 mm SL): rio das Fêmeas, near Barreiras, c. 12 ° 28 ’ S, 45 ° 13 ’ W; M. A. Cestarolli & J. Camargo, 2 – 6 May 1985. MZUSP 28771 (1, 140.8 mm SL): rio Desidério, São Desidério, 12 ° 21 ’ S, 44 ° 58 ’ W; M. A. Cestarolli & J. Camargo, 2 – 6 May 1985. MZUSP 70220 (4, 1 skel., 252.8 – 320.0 mm SL): Bahia, Barra, fish market, c. 11 ° 6 ' S, 43 ° 9 ’ W; O. T. Oyakawa et al., 10 April 2001. MZUSP 98750 (1, 92.2 mm SL): Barra, rio Grande, praia Cabeça de Touro, 11 ° 6 ’ 8 ’’ S, 43 ° 9 ’ 26 ’’ W; O. T. Oyakawa et al., 10 Apr 2001. NMW 62941 (2, 248.0 – 249.0 mm SL): rio São Francisco, Barra, 11 ° 5 ’ S, 43 ° 8 ’ W; F. Steindachner et al. (Austrian Expedition), March 1903. UMMZ 216373 (1, 106.7 mm SL): Barra, rio São Francisco, 11 ° 5 ’ S, 43 ° 8 ’ W; J. R. Bailey, 8 Apr 1942. MZUSP 18558 (1, 192.4 mm SL): Marcos, Remanso, Sobradinho reservoir, c. 9 ° 38 ’ S, 42 ° 4 ’ W; S. F. Santos, 20 Jul 1975. MZUSP 18666 (2, 303.3 – 324.8 mm SL): rio São Francisco, downstream Sobradinho reservoir, c. 9 ° 26 ’ S, 40 ° 47 ’ W; J. Dias, 27 Sept 1976. MZUSP 2007 (13, 1 cs, 132.3 – 225.0 mm SL); CAS 11822 (1, 243.0 mm SL): rio São Francisco, between Juazeiro and Barra; E. Garbe, 1908. FMNH 56814 (2, 182.0 – 212.0 mm SL): Juazeiro, rio São Francisco, 9 ° 24 ’ S, 40 ° 30 ’ W; J. D. Haseman, 28 Nov 1907. UMMZ 216349 (1, 85.3 mm SL): Juazeiro, small oxbow ponds on E (downstream edge of town), c. 9 ° 24 ’ S, 40 ° 30 ’ W; J. R. Bailey & J. M. de Oliveira, 18 Apr 1942. Pernambuco: UMMZ 147396 (2, 142.1 – 148.7 mm SL): Santa Maria da Boa Vista, “ Lake Aripos ”, c. 8 ° 48 ’ S, 39 ° 49 ’ W; R. S. Menezes, c. 1945. MZUSP 3797 (3, 83.5 – 135.1 mm SL): rio São Francisco (precise locality unknow); A. P. Marques, 1941. Locality uncertain (see Distribution): Piauí: MCZ 21268 (2, 152.3 – 167.2 mm SL): “ rio Puty, Therezina ” (rio Poti, tributary of rio Parnaíba at Teresina, 5 ° 5 ’ S, 42 ° 49 ’ W); O. St. John, Dec 1865.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F2EFF994EA4FEA8FE87FD17.taxon	description	S. (almo) matrincham Kner (ex Natterer), 1860: 12 (name not available, published as a synonym of Chalceus carpophagus Valenciennes).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F2EFF994EA4FEA8FE87FD17.taxon	diagnosis	Diagnosis. Brycon orbignyanus can be distinguished from all remaining cis-andean Brycon species, except for B. orthotaenia, B. hilarii, B. whitei, and B. polylepis, by possessing a caudal peduncle blotch extending as a stripe into the distal portion of caudal-fin rays (caudal peduncle blotch, when present, limited to the caudal peducle or extending only into centralmost caudal-fin rays; Fig. 5). It can be distinguished from these species by possessing dentary teeth decreasing gradualy in size (vs. four anteriormost teeth considerably larger than remaining teeth). It can be further distinguished from B. orthotaenia, B. hilarii, and B. whitei by possessing a relatively narrow and elongated dentigerous premaxilary surface (vs. a broad, relatively short dentigerous surface of the premaxillary). It can additionaly be distinguished from Brycon hilarii and B. whitei primarily by possessing lower scale counts (52 – 63, modally 56 lateral line scales, vs. 67 – 82, modally 74`in B. hilarii, and 66 – 76, modally 70, in B. whitei; 10 – 13, modally 12 scales between lateral line and dorsal-fin basis, vs. 12 – 17, modally 15, in B. hilarii, and 12 – 13, modally 13, in B. whitei; 19 – 23, modally 20 circumpeduncular scales, vs. 20 – 28, modally 26, in B. hilarii, and 19 – 24, modally 21, in B. whitei, respectively). Brycon orbignyanus can be further distinguished from B. whitei by lacking a midlateral dark stripe (vs. mdilateral dark stripe present). Brycon orbignyanus can be further distinguished from B. orthotaenia by possessing a pointed head profile (versus aproximately rounded to obtuse in the latter species; compare Figs. 6 F and H), by possessing a higher number of teeth in the second, inner premaxilary row (not counting the teeth of the second row situated between the first and third rows) (5 – 9, modally 6, versus 3 – 5, modally 5, in B. orthotaenia), and by possessing the dentary teeth decreasing gradualy in size (vs. anteriormost four dentary teeth considerably larger than the remaining teeth in B. orthotaenia).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F2EFF994EA4FEA8FE87FD17.taxon	description	Description. Morphometric data are presented in Table 15. Large-sized species, largest examined specimen 516.0 mm SL. Body moderately slender. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly concave to straight 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 to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Nine (3), 10 (8), 11 (15), 12 (18), or 13 (5) relatively small teeth in outer series. Teeth generally tricuspidate, some pentacuspidate in larger (> 470 mm SL) specimens. Five (8), 6 (24), 7 (17), 8 (2), or 9 (1) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 2 (7), 3 (35), or 4 (10) 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, penta- to hexacuspidate. Maxillary margins approximately parallel, straight in profile. Eleven to 20 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tri- to pentacuspidate, posterior teeth unicuspidate. Dentary with 8 (3), 9 (4), 10 (7), 11 (9), 12 (14), 13 (12), or 14 (5) teeth in main series. Teeth at dentary symphysis larger, pentacuspidate, remaining teeth decreasing gradually in size, larger teeth near dentary symphysis penta-, tetra-, tri- 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 12 – 15 small, aciculated, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of fifth to eighth main series dentary teeth. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Fifty-two (3), 53 (5), 54 (4), 55 (7), 56 (11), 57 (5), 58 (6), 59 (4), 60 (5), 61 (1), or 63 (1) 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 between 150 – 200 mm SL with tubules with two or three branches, four to seven branches in specimens between 220 – 270 mm SL, and with more than 10 branches and developing a dendritic pattern of ramification, with tubules overlapping each other in larger (> 300 mm SL) specimens. Horizontal scale rows between dorsal-fin origin and lateral line 10 (1), 11 (20), 12 (26), or 13 (4). Horizontal scale rows between lateral line and pelvic-fin 5 (4), 6 (18), 7 (24), 8 (4), or 9 (1). Circumpeduncular scales 19 (3), 20 (25), 21 (13), 22 (11), or 23 (1). Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 14 th (1) vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 21 (4), 22 (6), 23 (6), 24 (15), 25 (15), 26 (3), or 27 (2). First anal-fin pterygiophore inserting behind haemal spine of 28 th (1) 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 24 – 25 rectangular scales. Pectoral-fin rays i, 12 (4), 13 (40), 14 (8), or 15 (2). Pelvic-fin rays typically i, 7, a single specimen i, 6. Main caudal-fin rays 10 / 9. Caudal fin slightly forked, distal margin slightly concave. Central caudalfin rays 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 at the middle caudal-fin projection. 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 14 (3), 16 (1), 17 (15), 18 (12), 19 (6), or 20 (2) lower, 1 at angle, and 14 (1), 15 (4), 16 (4), 17 (16), 18 (10), or 19 (2) upper gill rakers. Vertebrae 47 (1), 49 (1). Supraneurals 11 (1). 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. 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, extending longitudinally to posterior margin of fifth 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. Caudal peduncle with broad median stripe, originating 4 – 6 scales from hypural joint and continuing posteriorly over 4 central principal caudal-fin rays to caudal-fin distal margin. Remaining caudal-fin rays, and remaining fins, clear. Coloration in life. Description based on photographs of freshly caught or living specimens from middle and upper rio Paraná in Brazil and Argentina (published pictures in Godoy, 1986: 76; Reis Filho, 1998: 54; Zaniboni Filho et al., 2004: 101). Overall coloration silvery, dorsum dark-grey, with a silvery hue. Top of head dark grey. Midlateral dark stripe extending across caudal peduncle and middle caudal-fin rays very conspicuous. Caudal-fin rays dorsal and ventral to middle dark stripe reddish to intense red. Anal, dorsal, and pelvic fins reddish. Pectoral fins orangish or dark. Sexual dimorphism. None of the examined specimens possess anal-fin hooks, but its presence was reported as a dimorphic feature of mature males in the species (Zaniboni Filho et al., 2004: 101). Common names. Brazil: “ piracanjuba ”, “ piracanjuva ” (Magalhães, 1931: 160; Godoy, 1975: 289); Argentina: “ pirapitá ”, “ salmón ”, “ salmón criollo ”, “ salmón del Paraná ”, “ salmonete ”, “ pracanjuva ”, “ pirapitanga ” (Ringuelet et al., 1967: 136), “ pirapitá blanco ” (F. Baena, pers. comm.); Uruguay: “ pirapitá ”, “ salmón ”, “ salmón criollo ”, “ pirapitanga ” (Sverlij et al., 1998: 16). Juvenile specimens were formerly called “ piracanjuvira ”, and adult, breeding specimens, “ piracanjuba arripiada ”, at the rio Mogi-Guaçú (Magalhães, 1931: 161; Godoy, 1975: 293).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F2EFF994EA4FEA8FE87FD17.taxon	distribution	Distribution. Originally widespread throughout the rio Paraná (both above and below the Sete Quedas waterfalls, the former barrier between the upper and lower sections of the rio Paraná) and rio Uruguai basins, in Brazil, Argentina, Paraguay, and Uruguay (Fig. 56; see also Liotta, 2005: 100, for a map of known localities in Argentina). It has now disappeared from most of its former range (see item “ Conservation ”, below). Brycon orbignyanus does not occur in the rio Paraguai basin, where it is replaced by the congener B. hilarii (see under item “ Distribution ” of this species), though both species occur syntopically in the middle rio Paraná in Brazil and Argentina. Ecological notes. Brycon orbignyanus originally occurred in middle to large sized rivers, and at floodplain lakes and channels associated with large rivers (Agostinho et al., 1997: 183). The species prefered forested rivers and was considered to be mainly frugivore (Magalhães, 1931; Ringuelet et al., 1967). Godoy (1975: 304 – 305) examined stomach contents of nine specimens collected at the rio Mogi-Guaçú and found vegetal matter (mostly leaves), insects, and small fishes. Hahn et al. (1997: 212) examined stomach contents of 46 specimens collected in the floodplains of the rio Paraná at Paraná state and, similarly to Godoy (1975), found insects and other invertebrates, vegetal matter, and fishes. Reis et al. (2003: 123) reported seeds but predominantly aquatic plants as the diet of six specimens collected in the upper rio Uruguai. Based on the findings by Hahn et al. (1997), Agostinho et al. (1997: 235) considered the species as being mainly insectivore, though that conclusion is probably mainly a result of the relative small size of the specimens examined by Hahn et al. (1997). Interestingly, the unique dentition of Brycon orbignyanus among congeners, with graduated-sized and relatively small and compressed anterior teeth on the dentary suggests a folivore diet for the species (pers. obs.). Size at first maturation is reported to be 30 cm TL, being reached at 2 – 3 years of age (Agostinho et al., 2003). Brycon orbignyanus is well-known as a migratory species. At the lower rio Paraná and rio Uruguai at Argentina and Uruguay, Brycon orbignyanus migrated downstream during October, moving upstream during March. Breeding in the southern part of the range of the species took place between December and January (Devincenzi & Teague, 1942: 73; Ringuelet et al., 1967: 137). On other hand, at the upper rio Paraná basin the reproductive migration is undertaken upstream (Ihering, 1929; Magalhães, 1931; Godoy, 1975). The upstream migration in rio Mogi-Guaçú started by the end of Setember, and spawning took place between November and January (Godoy, 1975: 296, 299). At the rio Piracicaba, Magalhães (1931: 162) reported reproductive schools moving upstream the river between December and January, and, on one occasion, hundreds of specimens were observed entering a flooded area, the males then apparently chasing the females. Ihering (1929: 81) estimated that females ranging between 570 and 690 mm in total length possessed a total fecundity of 500,000 to 1,000,000 oocytes. After spawning, Brycon orbignyanus move downstream, sometimes considerably far from the spawning site. Godoy (1975: 299 – 300) reported that two specimens captured and tagged at Cachoeira de Emas (rio Mogi Guaçú) were recaptured two to four months later 601 – 637 km downstream, into the rio Grande. Thormählen de Gil (1949: 414 – 437) studied annuli rings in scales and calculated that specimens between 90 – 116 mm SL are less than two years old, specimens between 123 – 215 mm SL are 2 – 3 years old and specimens between 229 – 245 mm SL are 4 – 5 years old. Thormählen de Gil (1949: 414 – 437) estimated that specimens with 264, 374, 530 and 604 mm SL were respectively about 6, 11, 13, and 16 years old. Though age estimation through annuli rings can be misleading as an age determinator (Casselman, 1990), so are other biological markers (Campana, 2001), and Thormählen de Gil’s (1949) data representa at the very least the only attempt to infer growth and longevity in wild populations of this now threatened fish. Godoy (1975: 294) reported that the largest female specimen recorded in the rio Mogi-Guaçú reached 79.5 cm TL and weighted 8.2 kg, while the largest male measured 68.0 cm TL and weighted 3.62 kg.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F2EFF994EA4FEA8FE87FD17.taxon	description	Conservation. Brycon orbignyanus was in the past one of the most valued and important fishes in commercial and sport fisheries throughout its range (e. g., Ringuelet et al., 1967; Schubart, 1943, 1949; Monteiro, 1953; Machado et al., 1968; Godoy, 1975). Godoy (1975) documented the gradual decline of the species in the rio Mogi- Guaçu in the upper rio Paraná basin, where the species was once one of most important target of fisheries (Schubart, 1943, 1949). The species is now extirpated from the rio Mogi-Guaçu, and, in fact, from almost the entire upper rio Paraná basin, with the exception of the last natural stretch of the rio Paraná in Brazil, the floodplains situated between the Porto Primavera and Itaipu dams (Agostinho et al., 2008). Populations of Brycon orbignyanus are in fact steeply declining throughout its entire range (Quirós, 1990; Agostinho et al., 2008; Mello et al., 2011). Though pollution and removal of the riparian forest were certainly important factors that contributed to the decline of the species, the disappearance of this previous widespread species throughout most of its range should be primarily imputed to the drastic alterations in flow regime caused by damming. The species clearly is unable to survive in reservoirs or river stretches with flow regimes regulated by hydroelectric dams. There is almost no freeflowing river stretches in the upper rio Paraná basin (Agostinho et al., 2003), and a similar fate is under way for the upper rio Uruguai basin. Brycon orbignyanus is officially considered a threatened species in both Brazil (Agostinho et al., 2008) and Argentina, and should in fact be considered as globally threatened. There is a relatively intense stocking program for the species in Brazil (e. g., Dumont-Neto et al., 1997; Senhorini et al., 2002; Lopera et al.. 2008).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F2EFF994EA4FEA8FE87FD17.taxon	discussion	Remarks. Chalceus orbignyanus was described by Valenciennes (in Cuvier & Valenciennes, 1850: 249), based on two specimens (MNHN 9835) collected at “ Buénos-Ayres ”, collected by Alcides d’Orbigny. Géry & Mahnert (1992: 811) designated the larger syntype of the lot MNHN 9835 (161 mm SL) as the lectotype. As defined in the present study, Brycon orbignyanus possess three junior synonyms. Chalceus rodopterus Valenciennes (in Cuvier & Valenciennes, 1850: 249 – 250), described subsequently to Chalceus orbignyanus based on two juvenile specimens also collected at “ Buénos-Ayres ”, is, as already remarked by Géry & Mahnert (1992: 807 – 808), a synonym of B. orbignyanus. Brycon lineatus, described by Steindachner (1866 a: 19) and redescribed and illustrated by the same author (Steindachner, 1866 b: 211 – 212, pl. 2), was also considered by Géry & Mahnert (1992: 807) as a possible synonym of B. orbignyanus, differing, however, by possessing a lower number of precaudal vertebrae (21) and supraneurals (8). Those counts are indeed lower than the usual counts for the species (28 to 29 pre-caudal vertebrae, and 11 – 12 supraneurals). Géry & Mahnert (1992), which have only examined the radiograph of the holotype of Brycon lineatus, supposed that it could be a teratological specimen. The examination of the purported holotype of Brycon lineatus (NMW 62943) revealed it to be rather a specimen of Brycon falcatus, a species actually quite distinct from B. orbignyanus, which explains the discrepancy in pre-caudal vertebrae and supraneurals noticed by Géry & Mahnert (1992). Since the description and illustration by Steindachner (1866 b) clearly depicts a specimen of Brycon orbignyanus, which is additionaly the sole species of Brycon occurring in the southern portion of the La Plata basin, it seems that at some point the holotype of Brycon lineatus was inadvertently exchanged with this Brycon falcatus specimen. Although the “ true ” holotype of Brycon lineatus is apparently lost, we concur with Géry & Mahnert (1992) in concluding that this nominal species clearly represents a synonym of B. orbignyanus.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F2EFF994EA4FEA8FE87FD17.taxon	description	Some authors (Magalhães, 1931; Schubart, 1943, 1962; Godoy, 1975) erroneously identified B. orbignyanus from the upper rio Paraná basin in São Paulo (and, particularly, from the rio Mogi Guaçú basin) as Triurobrycon lundii (= Brycon orthotaenia). All specimens examined in the present study from the rio Mogi-Guaçu basin (MZUSP 2070, MZUSP 3830, BMNH 1946.12.23.136 - 137) are undoubtedly Brycon orbignyanus, B. orthotaenia being restricted to the rio São Francisco basin (see under this species).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F2EFF994EA4FEA8FE87FD17.taxon	materials_examined	Material examined. Type material: MNHN 9835 (2, 142.7 – 160.9 mm SL): “ Buenos-Ayres ”; A. d’Orbigny. Lectotype (larger specimen) and paralectotype (smaller specimen) of Chalceus orbignyanus Valenciennes, 1850 (see Géry & Mahnert, 807, 811). MNHN A. 9834 (2, 78.8 – 79.5 mm SL): “ Buenos-Ayres ”; A. d’Orbigny. Syntypes of Chalceus rodopterus Valenciennes, 1850. MZUSP 3811 (1, 241.0 mm SL): Mato Grosso [= Mato Grosso do Sul?], rio Bodoquena [=? Bodoquena; see “ Remarks ”, above]; L. Travassos-Filho, 1941. Holotype of Brycon travassosi Amaral Campos, 1950. Non types. Brazil, Goiás: NUP 1125 (2, 223.9 – 240.9 mm SL): Caldas Novas, rio Corumbá (trib. rio Paranaíba) at Corumbá reservoir, 17 ° 43 ’ 37 ’’ S, 48 ° 32 ’ 54 ’’ W; Nupélia team, 9 Apr 1999. São Paulo: MZUSP 16626 (1, 280.8 mm SL): Rio Grande, Cachoeira do Onça (now under Ilha Solteira dam), 19 ° 59 ’ S, 50 ° 53 ’ W; P. E. Vanzolini et al., 6 – 7 Jan 1962. MZUSP 17169 (7, 200.8 – 473.8 mm SL): Ilha Solteira, rio Paraná, 20 ° 23 ’ S, 51 ° 21 ’ W; Exc. Departamento de Zoologia, Sept 1965. MZUSP 67070 (1, 156.5 mm SL): rio Aguapeí (trib. rio Paraná), c. 21 ° 03 ’ S, 51 ° 46 ’ W; J. Canella, 27 Aug 1941. MZUSP 67069 (1, 167.1 mm SL): rio Paraná, mouth of rio Aguapeí, 21 ° 03 ’ S, 51 ° 46 ’ W; J. Canella, 7 Aug 1941. MNRJ 4776 (1, 209.0 mm SL); MNRJ 4767 (1, 187.8 mm SL); MNRJ 4775 (1, 198.2 mm SL); MNRJ 4779 (12, 99.6 – 289.0 mm SL): rio Paraná, Porto Cabral, 22 ° 17 ’ S, 52 ° 38 ’ W; L. Travassos Filho, March – April 1944. FMNH 119092 (1, 200.0 mm SL): rio Tietê, Salto do Avanhandava, c. 21 ° 13 ’ S, 49 ° 57 ’ W; J. D. Haseman, 15 Sept 1908. FMNH 92082 (1, 213.0 mm SL): Itapura, rio Tietê, 20 ° 39 ’ S, 51 ° 30 ’ W; J. D. Haseman, 27 Sept 1908. MZUSP 1474 (1, 147.9 mm SL): Itapura, rio Tietê, 20 ° 39 ’ S, 51 ° 30 ’ W; E. Garbe, 1904. MZUSP 79319 (1, 89.9 mm SL): Pereira Barreto, rio Tietê, Três Irmãos reservoir, 20 ° 40 ’ 25 ’’ S, 51 ° 30 ’ 6 ’’ W; W. S. Smith, Jan 2001. MZUSP 1523 (1, 171.9 mm SL): Piracicaba, rio Piracicaba, 22 ° 43 ’ S, 47 ° 40 ’ W; J. Lima, 1896. MZUSP 3900 (1 head and scapular girdle; 108.9 mm HL): Pirassununga, rio Mogi-Guaçú, Cachoeira de Emas, 21 ° 55 ’ S, 47 ° 22 ’ W; R. von Ihering, 1907. MZUSP 3830 (7, 115.3 – 195.0 mm SL): same locality; O. Schubart, 1945. BMNH 1946.12.23.136 - 137 (2, 120.4 – 234.0 mm SL): Rio Mogi-Guaçu (no precise locality but very likely Cachoeira de Emas); A. Amaral Campos, no date. MZUSP 3618 (1, 334.9 mm SL): same locality; A. Amaral Campos, March 1943. MZUSP 3378 (10, 2 cs, 109.9 – 129.4 mm SL): Teodoro Sampaio, rio Paranapanema at Porto Marcondes, 22 ° 32 ’ S, 52 ° 2 ’ W; E. Dente & D. Seraglia, 17 – 23 Nov 1946. Mato Grosso do Sul: MZUSP 18377 (1, 179.0 mm SL): Ilha Solteira, rio Paraná, opposite to Ilha Solteira (cofferdam), 20 ° 23 ’ S, 51 ° 21 ’ W; Exc. MZUSP, 25 – 28 May 1972. MZUSP 16714 (3, 211.3 – 277.4 mm SL): Três Lagoas, rio Paraná, at Jupiá, 20 ° 47 ’ S, 51 ° 38 ’ W; Exc. Departamento de Zoologia, 15 – 23 Sept 1962. MZUSP 3039 (1, 213.8 mm SL): rio Verde (trib. rio Paraná), 21 ° 11 ' S 51 ° 53 ' W; J. Canella, 25 Aug 1941. Paraná: MZUEL 7440 (34, 107.1 – 160.7 mm SL); ZUEC 7762 (4, 121.6 – 146.9 mm SL): Icaraíma, rio Paraná, Ilha Maringá, c. 23 ° 21 ’ S, 53 ° 47 ’ W; O. A. Shibatta & M. Caetano Filho, 31 March – 2 April 1995. MZUEL 5387 (2, 188.0 – 240.0 mm SL): Altônia, rio Paraná, Porto Cerâmica, 23 ° 51 ’ 40 ’’ S, 50 ° 0 ’ 48 ’’ W; F. Yuldi, 25 March 2010. LBP 9210 (1, 190.0 mm SL): Altônia, rio Paraná, Porto Cerâmica, 23 ° 51 ’ 40 ’’ S, 54 ° 0 ’ 48 ’’ W; A. C. R. Casimiro & F. Yuldi, 25 Apr 2010.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F2EFF994EA4FEA8FE87FD17.taxon	description	MZUSP 42353 (2, 179.0 – 196.3 mm SL): Reservatório de Itaipu, c. 25 ° 16 ’ S, 54 ° 28 ’ W; no collector specified, Jan 1990. MZUSP 18840 (1, 237.7 mm SL): Guaíra, rio Paraná, 24 ° 4 ’ S, 54 ° 15 ’ W; CETESB, Jul – Aug 1977. MZUSP 21065 (2, 195.0 – 204.2 mm SL): Guaíra, rio Paraná below Sete Quedas waterfalls, c. 24 ° 7 ’ S, 54 ° 20 ’ W; CETESB, 1977 – 1980. MZUSP 15364 (1, 443.6 mm SL): Porto Mendes, rio Paraná, 24 ° 29 ’ S, 54 ° 19 ’ W; CETESB, Nov – Dec 1977. Santa Catarina: MCP 21605 (1, 322.7 mm SL): Itapiranga, rio Uruguai, near pedra da Fortaleza, 27 ° 11 ' 42 '' S 53 ° 38 ' 34 '' W; L. F. Câmara et al., 5 Nov 1998. MCP 22458 (1, 373.5 mm SL); MCP 22457 (1, 326.1 mm SL): same locality; L. Hahn et al., 26 Feb 1999. Rio Grande do Sul: MCP 8746 (1, 516.0 mm SL); MCP 8747 (1, 506.5 mm SL); rio Uruguai (no specific locality); J. Bertoletti, June 1975. Paraguay: UMMZ 207060 (1, 515.0 mm SL): Pettirossi fish market (= Mercado Quatro) in Asuncion; J. Taylor et al., 14 Sept 1979. Argentina: MZUSP 17115 (1, 111.1 mm SL): Rio de la Plata, Buenos Aires, c. 34 ° 30 ’ S, 58 ° 26 ’ W; H. P. Castello, 4 April 1965. BMNH 1927.2.9.7 - 9 (4, 188.9 – 205.3 mm SL): Misiones, Puerto Iguazu, rio Paraná at mouth of rio Iguazu, 25 ° 35 ’ S, 54 ° 35 ’ W; G. Harrison, no date. BMNH 1872.6. 8.22 (1, 307.0 mm SL): “ Paraná ” (no precise locality); collector and date not specified. Uruguay: MNRJ 11266 (1, 124.8 mm SL): Departamento Colonia, Arroyo Linetas, Estancia San Jorge, c. 34 ° 10 ’ S, 58 ° 6 ’ W; R. Pradori, 17 Apr 1965. MCZ 845 (1, 309.6 mm SL): Rio Uruguay (no precise locality); J. Wyman, no date.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F16FF934EA4FC83FF2CFF1F.taxon	description	Salmo para-putanga Kner (ex Natterer), 1860: 4 (published as a synonym of Chalceus orbygnianus; not available). Brycon melanoxanthus Kner (ex Natterer), 1860: 4 (published as a synonym of Chalceus orbygnianus; not available).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F16FF934EA4FC83FF2CFF1F.taxon	diagnosis	Diagnosis. Brycon hilarii can be distinguished from all remaining cis-andean Brycon, except for B. orthotaenia, B. orbignyanus, B. whitei, and B. polylepis by 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 polylepis by 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 orbignyanus and B. orthotaenia primarily by possessing higher scale counts (67 – 82, modally 74, vs. 52 – 63, modally 56, and 49 – 58, modally 52 lateral line scales in Brycon orbignyanus and 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. orbignyanus and B. orthotaenia, respectively; 20 – 28, modally 26, vs. 19 – 23, modally 20, and 18 – 21, modally 19 circumpeduncular scales in B. orbignyanus and B. orthotaenia, respectively). Brycon hilarii can be distinguished from B. whitei by 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. hilarii and the latter species in the Peruvian Amazon.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F16FF934EA4FC83FF2CFF1F.taxon	description	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 mm SL, 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. 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 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 four branches in specimens between 150 – 230 mm SL, and with more than 10 branches and developing a dendritic pattern of ramification, with tubules overlapping each other in larger (> 300 mm SL) 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 13 th (6) to 14 th (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 25 th (4) or 26 th (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. Central caudalfin 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 mm SL) 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. 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 Plata system 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 Plata system 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 type of 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).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F16FF934EA4FC83FF2CFF1F.taxon	distribution	Distribution. Rio Paraguai and middle rio Paraná basins in Brazil and Paraguay, and upper Rio Amazonas basin in Peru, Ecuador, and Colombia (Fig. 62). Brycon hilarii is recorded for the rio Paraná below 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 Paraná between 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 hilarii also 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 hilarii from upper tributaries of the Amazon basin in Colombia, but a photograph of a specimen collected at the upper Río Putumayo provided by A. Linares confirm the occurrence of the species in that country. Although Géry & Mahnert (1992: 800) record Brycon hilarii from the upper rio Madeira system in Bolivia, we have only examined specimens of B. amazonicus from this latter area and consider that the eventual occurrence of the species at the upper Rio Madeira basin in Bolivia needs further corroboration. The species is being extensively stocked in eastern Brazil and accidental / intentional releases in the upper rio Paraná basin are reported (Graça & Pavanelli, 2007: 77). Ecological notes. Brycon hilarii is a relatively eurytopic species in the rio Paraguai basin in Brazil and Paraguay, occurring from the large rivers in the lowlands and associated floodplains to relatively small, fastflowing rivers in the headwaters. At the middle rio Paraná basin in Brazil and Argentina Brycon hilarii occurs syntopically with B. orbignyanus. The largest specimen recorded in the literature measured 670 mm TL (Mateus & Estupiñan, 2002: 166). Diet and feeding behavior in the rio Paraguai basin in Brazil were studied by Sabino & Sazima (1999) and Reys et al. (2009). As other Brycon species, B. hilarii is 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 hilarii in the rio Formoso (Reys et al., 2009). Reys et al. (2009) hypothesized that Brycon hilarii acted 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 hilarii consisting in up to 30 specimens following capuchin monkeys (Cebus apella) to catch fruits (specifically, Guarea cf. 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 13 C 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 hilarii undertake 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 Miranda in Mato Grosso do Sul state is said to migrate downstream into the turbid, relatively warm rio Miranda to spawn (Resende, 2011). The species is one of most valued and important targets of both commercial and recreational fisheries of the rio Paraguai basin 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. hilarii from the rio Miranda by 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.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F16FF934EA4FC83FF2CFF1F.taxon	discussion	Remarks. Valenciennes (in Cuvier & Valenciennes, 1850: 247 – 248) described Chalceus hilarii based 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 hilarii showed that these specimens clearly are not conspecific with the lectotype but rather are referrable to B. falcatus. After its original description, Brycon hilarii was 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 Brycon species occurring in the rio Paraguai basin. They considered that the name Chalceus hilarii could 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 hilarii might be a senior synonym of Brycon orthotania from the rio São Francisco. This change in name usage, from Brycon hilarii to 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 hilarii for the species from the rio Paraguai, and considered B. microlepis to 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. orthotaenia showed 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 hilarii is much more similar to Brycon specimens from the rio Paraguai basin, than to the species occurring at the rio São Francisco basin. We thus consider that the name Chalceus hilarii should in fact be applied to the Brycon species 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 hilarii was 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 Spatuloricaria is known from the rio Paraguai basin. The systematics of Spatuloricaria are 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 nudiventris corresponds to the Spatuloricaria species 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 hilarii and the holotype of Loricaria nudiventris were 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 erythropterum was distinguished from the species then assigned by Cope to the genus Megalobrycon (M. cephalus and 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 hilarii populations from the upper Amazon basin, and, consequently, Megalobrycon erythropterum is herein considered as a synonym of the latter species.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F16FF934EA4FC83FF2CFF1F.taxon	materials_examined	Material examined. Type material: MNHN A. 8616 (1, 423.3 mm SL): “ Rio San Francisco ” [type-locality very probably incorrect, see Discussion, above]; A. Saint-Hilaire, no date. Lectotype of Chalceus hilarii Valenciennes, designated by Géry & Mahnert (1992: 815). Non types. Brasil, rio Paraguai basin. Mato Grosso: MZUSP 81773 (1, 267.4 mm SL): Nobres, rio Manso basin, c. 14 ° 45 ’ S, 56 ° 19 ’ W; J. M. Mendes, no date. MZUSP 81126 (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. MZUSP 90089 (1, 103.5 mm SL): 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. MZUSP 18651 (1, 252.0 mm SL): Cáceres, rio Paraguai, 16 ° 4 ’ S, 57 ° 41 ’ W; CEPIPAM, 17 – 19 Feb 1976. CAS 18358 (1, 244.0 mm SL): rio Paraguai, São Luiz de Cáceres, 16 ° 4 ’ S, 57 ° 41 ’ W; J. D. Haseman, 27 May 1909. MCP 15817 (2, 144.8 – 184.2 mm SL): Cáceres, rio Paraguai at Cáceres and surroundings, 16 ° 4 ’ S, 57 ° 41 ’ W; R. E. Reis et al., 11 Aug 1991. MCP 10752 (1, 174.3 mm SL): rio Pixaim, Transpantaneira road, 60 km south 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 Transpantaneira road, c. 16 ° 26 ’ S, 56 ° 40 ’ W; J. C. Garavello et al., 16 – 22 April 1981. ZUEC 6853 (2, 128.8 – 157.7 mm SL): Poconé, Transpantaneira road, km 10, c. 16 ° 28 ’ S, 56 ° 41 ’ W; I. Sazima & F. A. Machado, 27 Apr 1981. ZUEC 3207 (1, 107.3 mm SL): Poconé, rio Piraputanga, fazenda Ipiranga, c. 16 ° 30 ’ S, 56 ° 45 ’ W; F. A. Machado et al., 16 May 1997. MZUSP 19240 (1, 193.9 mm SL): Poconé, c. 16 ° 29 ’ S, 56 ° 41 ’ W; G. Schaller, 1978. MNRJ 20525 (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. MZUSP 4379 (1, 151.5 mm SL): Santo Antônio do Leverger, rio Cuiabá, 15 ° 52 ’ S, 56 ° 4 ’ W; G. Olson, 1965. MZUSP 95016 (1, 176.2 mm SL): 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. MZUSP 89522 (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. ZUEC 5705 (1, 130.7 mm SL): Santo Antônio de Leverger, flooded areas near Rio Cuiabá, c. 15 ° 53 ’ S, 56 ° 2 ’ W; F. A. Machado et al., 28 May 1989. MZUSP 85579 (8, 3 cs, 71.2 – 124.3 mm SL): Barão de Melgaço, flooded areas near Mimoso, c. 16 ° 17 ’ S, 55 ° 49 ’ W; F. A. Machado, 19 Jan 1999. MZUSP 18795 (8, 273.2 – 304.4 mm SL): Barão de Melgaço, rio Cuiabá, 16 ° 11 ’ S, 55 ° 58 ’ W; CEPIPAM, 1 – 10 May 1977. INPA 16369 (1, 161.9 mm SL): Baía Sinhá Mariana, c. 30 km below Barão de Melgaço, c. 16 ° 20 ’ S, 55 ° 54 ’ W; CEPIPAM, 30 Nov 1979. MZUSP 67189 (71, 5 cs, 23.4 – 45.3 mm SL): Barão de Melgaço, flooded areas at Fazenda do Poli; K. de Silimon, 19 Dec 1983. ZUEC 5680 (1, 149.9 mm SL): Cuiabá, rio Cuiabá; F. A. Machado, 2 May 1981. FMNH 76445 (1, 196.0 mm SL): Rio Paraguai, Descalvados, 16 ° 43 ’ S, 57 ° 45 ’ W; Schmidt & Sanborn, 20 Sept 1926. MZUSP 27178 (2, 192.6 – 199.9 mm SL): rio Paraguai, ilha de Taiamã, c. 16 ° 50 ’ S, 57 ° 34 ’ W; A. S. Soares, 1 – 7 Dec 1980. MZUSP 35871 (1, 190.7 mm SL): Itiquira, Baía Grande, fazenda Santo Antônio do Paraíso, c. 17 ° 33 ’ S, 55 ° 16 ’ W; J. H. B. Medeiros & J. C. Oliveira, 30 April 1979. MZUSP 25284 (1, 180.3 mm SL): Itiquira, Baía Grande, fazenda Santo Antônio do Paraíso, c. 17 ° 33 ’ S, 55 ° 16 ’ W; J. C. Oliveira, 24 – 29 Oct 1978. Mato Grosso do Sul: MZUSP 18681 (1, 243.4 mm SL); MZUSP 38206 (1, 364.0 mm SL): Coxim, rio Taquari, Cachoeira das Palmeiras, 18 ° 21 ’ 45 ’’ S, 56 ° 36 ’ 45 ’’ W; CEPIPAM, 6 – 8 Dec 1976. MZUSP 18556 (1, 170.7 mm SL): Coxim, rio Taquari, c. 18 ° 29 ’ S, 54 ° 45 ’ W; A. Storti & W. Uieda, 15 Aug 1975. MZUSP 2910 (5, 231.7 – 268.1 mm SL): Coxim, rio Piquerí, c. 17 ° 55 ’ S, 54 ° 41 ’ W; J. Lima, Oct 1930. MZUSP 17299 (1, 215.1 mm SL): rio Taquari, c. 150 km from Coxim; J. C. Garavello et al., 9 – 22 Aug 1967. BMNH 1910.5.26.19 – 20 (2, 244.0 – 284.0 mm SL): “ Pan de Ezucar, R. Paraguay ” (not located); C. Grant, no date. MZUSP 48297 (2, 1 cs, 154.3 – 161.2 mm SL): Fazenda Santo Antônio (Baía da Sede), Pantanal de Paiaguás, c. 17 ° 53 ’ S, 57 ° 8 ’ W; T. Liparelli, no date. MZUSP 36444 (1, 171.9 mm SL): Corumbá, Corixão, Capão Grande, Nhecolândia; G. M. Mourão et al., Sept 1985. MZUSP 18936 (1, 247.8 mm SL): rio Paraguai, surroundings of Corumbá, 19 ° 00 ’ S, 57 ° 38 ’ W; CEPIPAM, 12 Oct 1977. CAS 18359 (1, 257.8 mm SL), lagoons and rivers near Corumbá, 19 ° 0 ’ S, 57 ° 38 ’ W; J. D. Haseman, 28 April 1909. MZUSP 63173 (1, 221.8 mm SL): Corumbá, rio Paraguai, Baía de Albuquerque, 19 ° 24 ’ 54 ’’ S, 57 ° 22 ’ 43 ’’ W; CEPIPAM, 19 Feb 1977. MZUSP 49023 (3, 13.8 – 18.4 mm SL): Corumbá, Passo do Lontra, 19 ° 34 ’ S, 57 ° 1 ’ W; Exp. Zool. UFMS, 13 Jan 1988. LBP 37 (3, 141.3 – 151.9 mm SL): 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); MZUSP 59527 (1, 181.6 mm SL): Corumbá, rio Abobral 2, 19 ° 26 ’ 68 ’’ S, 57 ° 02 ’ 49 ’’ W; A. Machado-Allison et al., 7 Sept 1998. MZUSP 59528 (1, 133.2 mm SL): Aquidauana, fazenda São Pedro, 19 ° 36 ’ 70 ’’ S, 56 ° 02 ’ 78 ’’ W; A. Machado-Allison et al., 4 Sept 1998. MZUSP 2986 (1, 232.3 mm SL): Miranda, Salobra, rio Miranda, 20 ° 12 ’ S, 56 ° 30 ’ W; L. Travassos, Aug 1940. MNRJ 2878 (1, 286.0 mm SL): Miranda, rio Miranda, Salobra, 20 ° 12 ’ S, 56 ° 30 ’ W; L. Travassos, 1941. MZUSP 3073 (1, 183.7 mm SL): “ rio Bodoquena ” (precise locality uncertain, see under Remarks of Brycon orbignyanus); L. Travassos, 1941. Paraná, rio Paraná basin: MZUSP 21064 (7, 277.4 – 357.9 mm SL): Guaíra, rio Paraná below Sete Quedas waterfalls, c. 24 ° 7 ’ S, 54 ° 20 ’ W; CETESB, 1977 – 1978. MZUSP 14652 (1, 374.0 mm SL); MZUSP 14653 (1, 327.0 mm SL); MZUSP 14654 (1, 330.0 mm SL): Porto Mendes, rio Paraná, 24 ° 29 ’ S, 54 ° 19 ’ W; CETESB, 1978. Paraguay, rio Paraguai basin: FMNH 108173 (1, 262.0 mm SL): Depto. Alto Paraguay, Rio Negro, ca. 2.5 km above mouth of the Rio Paraguay, 20 ° 9 ' S, 58 ° 10 ' W; D. Mandelburger et al., 5 Sept 1997. FMNH 108174 (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. USNM 232317 (1, 257.1 mm SL): Depto. Amambay, Parque Nacional Cerro Cora, Rio Aquidaban, c. 22 ° 38 ’ S, 56 ° 1 ’ W; L. Naylor & C. Cuevas, 16 Jan 1982. UMMZ 206816 (1, 233.8 mm SL): Depto. Concepción, Arroyo Cagata, c. 14 km S of Yby-Yaú, 23 ° 6 ’ 30 ’’ S, 56 ° 31 ’ 12 ’’ W; J. Taylor et al., 28 Jul 1979. CAS 77377 (1, 174.0 mm SL): Depto. Concepción, Arroyo Trementina, trib. of the R. Aquido Canigi (= Rio Aquidaban?), c. 23 ° 3 ’ S, 57 ° 0 ’ W; J. D. Anisits, Dec 1900. USNM 181763 (1, 247.7 mm SL): Depto. Central, Asuncion bay, Río Paraguay near Asuncion, c. 25 ° 16 ’ S, 57 ° 38 ’ W; C. J. D. Brown, 15 Jan 1957. UMMZ 208068 (1, 278.8 mm SL): Depto. Central, Río Paraguay, overflow c. 1 km below Puerto Remanzo bridge, c. 25 ° 11 ’ S, 57 ° 33 ’ W; J. Taylor et al., 15 Sept 1979. USNM 181762 (2, 236.2 – 239.0 mm SL): Depto. Cordillera, Arroyo Pirareta, near Piribebuy, c. 25 ° 30 ’ S, 56 ° 56 ’ W; C. J. D. Brown, 13 Dec 1956. UMMZ 205665 (1, 237.1 mm SL): Depto. Cordillera, Salto de Pirareta, c. 400 m below fall, c. 25 ° 30 ’ S, 56 ° 56 ’ W; J. N. Taylor et al., 25 – 26 May 1979. USNM 181761 (1, 234.3 mm SL): Río Tebicuary near “ Reciefe ” (not located); C. J. D. Brown, 6 Dec 1956.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F16FF934EA4FC83FF2CFF1F.taxon	description	Rio Amazonas basin, Peru. Depto. Loreto: MZUSP 26507 (1, 228.7 mm SL): Río Ucayali, “ Albufao ” (not located); J. Guevara, 9 Oct 1971. NRM 23675 (1, 230.3 mm SL): El Estrecho, Río Putumayo, 2 ° 27 ’ S, 72 ° 40 ’ W; S. O. Kullander et al., 18 Jul 1986. MUSM 12745 (1, 132.6 mm SL): Yanayacu, Base 3, Pluripetrol, Río Shiriyacu, 4 ° 51 ’ S, 74 ° 56 ’ W; H. Ortega, 10 Oct 1997. Depto. Ucayali: MZUSP 26064 (1, 213.6 mm SL); MUSM 31 (190.0 mm SL); MUSM 30 (1, 215.0 mm SL): Río Huacamayo, road Pucallpa-Huánuco (= carretera Aguaytia / Pucallpa), km 7.5; J. Guevara, 17 Oct 1971. MUSM uncat. (1, 190.9 mm SL): Pucallpa, Río Neshuya, IVITA, 8 ° 38 ’ S, 74 ° 58 ’ W; H. Ortega, 6 Dec 1971. MZUSP 26471 (1, 83.2 mm SL): Río Neshuya, road Pucallpa-Huánuco, 8 ° 38 ’ S, 74 ° 58 ’ W; H. Ortega, 23 Jan 1976. MZUSP 26373 (1, 108.8 mm SL): Río San Alejandro, Pucallpa, c. 8 ° 50 ’ S, 75 ° 13 ’ W; J. Guevara, 14 Nov 1971. MUSM 28 (1, 248.0 mm SL): Río Saballo, trib. Río Aguaytia, Aguaytia, Tangarana; J. Guevara, 14 Oct 1971. MUSM 28573 (1, 167.4 mm SL): Padre Abad, Río Aguaytia basin, Río Santa Ana, Quebrada Samíria, 8 ° 32 ’ 41 ’’ S, 75 ° 38 ’ 35 ’’ W; M. Hidalgo & M. Rojas, 5 Dec 2006. MUSM 31 (2, 248.0 – 288.1 mm SL): Porto of Pucallpa, Río Ucayali; H. Ortega, 10 Oct 1971. MUSM 381 (1, 159,0 mm SL): Pucallpa, Tachsitea, Río Calleria (trib. Río Ucayali), 8 ° 1 ’ S, 74 ° 35 ’ W; P. de Rham & H. Ortega, 4 Oct 1984. MUSM 23 (1, 165.0 mm SL): Pucallpa, Río Ucayali, Masisea, 8 ° 35 ’ S, 74 ° 20 ’ W; H. Ortega, 24 Sept 1975. MUSM 41413 (1, 79.1 mm SL): Pucallpa, Río Ucayali, 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. FMNH 84312 (1, 267.0 mm SL): mouth of Rio San Alejandro at junction with Sungaro Yacu, c. 8 ° 27 ’ S, 75 ° 11 ’ W; G. Gutierrez, 1 Aug 1975. FMNH 84095 (1, 393.0 mm SL): Rio San Alejandro, deep pool upstream from camp, c. 8 ° 27 ’ S, 75 ° 11 ’ W; G. S. Glodek, 3 Aug 1975. FMNH 84094 (1, 383.0 mm SL): Rio San Alejandro, c. 8 ° 27 ’ S, 75 ° 11 ’ W; D. W. Greenfield, 3 Aug 1975. CAS 16025 (1, 275.1 mm SL); CAS 68879 (ex IU 16025) (2, 162.7 – 166.9 mm SL): Lago Cashiboya, a cutoff lake of Río Ucayali (connected to river by a channel) above Contamana, 7 ° 40 ’ S, 74 ° 56 ’ W; W. R. Allen, 3 – 4 Aug 1920. CAS 68878 (ex IU 16023) (3, 151.8 – 186.0 mm SL): Río Ucayali at Orellana, 6 ° 55 ’ S, 75 ° 9 ’ W; W. R. Allen, 9 Aug 1920. Depto. Huánuco: NRM 23669 (1, 150.2 mm SL): near mouth of quebrada trib. Río Pachitea, right bank, 2 – 3 km downstream Tournavista, c. 8 ° 57 ’ S, 74 ° 41 ’ W; S. O. Kullander & A. Hogeborn, 19 Aug 1981. CAS 68881 (ex IU 16026) (1, 196.2 mm SL): Río Pachitea; W. R. Allen, 20 – 22 Aug 1920. ROM 55913 (1, 167.3 mm SL): Río Llullapichis (trib. Río Pachitea), Panguana, Río Pachitea drainage, c. 9 ° 37 ' S, 74 ° 56 ' W; H. Pandura, Jul 1988. Ecuador: FMNH 103396 (1, 322.2 mm SL): Napo, Tiuyacu, first tributary to Rio Churuayacu upstream from mouth at Rio Payamino and near mouth of Rio Churuyacu, 0 ° 29 ' 30 '' S, 77 ° 18 ' W; D. Stewart, M. Ibarra & R. Barriga, 13 Nov. 1983. MCZ 60933 (1, 298.0 mm SL): Napo, Río Punino, trib. Río Payamino, above Coca, c. 0 ° 30 ’ S, 77 ° 0 ’ W; T. R. Roberts & H. Pauker, 25 Nov 1971. FMNH uncat. (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.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F1CFF974EA4FE8BFC1CFA9F.taxon	diagnosis	Diagnosis. Brycon whitei can be unambiguously diagnosed from all cis-andean congeners by the possession of an autapomorphic broad, dark, midlateral stripe extending from supracleithrum area to caudal peduncle and into median caudal-fin rays. Among congeners, Brycon whitei is more similar to B. hilarii. Brycon whitei specimens with poorly developed longitudinal stripes are very similar to B. hilarii specimens with well-developed caudal peduncle stripes, which in those specimens extend forward to the level of dorsal fin. However, Brycon whitei can be additionaly diagnosed from the latter species by possessing tiny stripes on caudal-fin rays (vs. tiny stripes on caudal-fin rays absent in B. hilarii).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F1CFF974EA4FE8BFC1CFA9F.taxon	description	Description. Morphometric data are presented in Table 17. Large-sized species, largest examined specimen 343.5 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 concave 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 middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Eight (4), 9 (5), 10 (5), 11 (4), 12 (2), or 13 (1) relatively small tricuspidate teeth in outer series. Three (3), 4 (8), or 5 (10) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 3 (2), 4 (18), or 5 (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, both hexacuspidate. Maxillary margins approximately parallel, straight in profile. Twelve to 23 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 8 (2), 9 (2), or 11 (2) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, tetra- to pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, tri- 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 14 small, aciculated, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of fifth to sixth main series dentary teeth. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Sixty-six (4), 67 (3), 68 (3), 70 (7), 71 (1), 73 (1), 74 (1), or 76 (1) scales in lateral line series. Laterosensory tube ramified tubules ramification increasing in complexity along ontogeny, specimens up to 150 mm SL with tubules with two to four branches, three to five branches in specimens around 200 mm SL, and with more than 6 branches and developing a dendritic pattern of ramification, with tubules often overlapping each other in specimens around 250 mm SL. Horizontal scale rows between dorsal-fin origin and lateral line 12 (7) or 13 (14). Horizontal scale rows between lateral line and pelvic-fin 7 (7), 8 (9), or 9 (5). Circumpeduncular scales 19 (1), 21 (9), 22 (7), 23 (3), or 24 (1). Dorsal-fin rays ii, 9. Dorsal fin origin slightly after to slightly ahead middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 14 th (1) vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in cs specimens), 21 (7), 22 (7), 23 (3), or 24 (4). First anal-fin pterygiophore inserting behind haemal spine of 31 th (1) 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 – 21 rectangular scales. Pectoral-fin rays i, 12 (2), 13 (10), or 14 (9). Pelvic-fin rays typically i, 7, one specimen i, 6, and two specimens i, 8. Main caudal-fin rays 10 / 9. Caudal fin slightly forked, distal margin slightly concave. Central caudal-fin 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 15 (1) or 17 (1) lower, 1 at angle, and 13 (1) or 16 (1) upper gill rakers. Vertebrae 47 (1). Supraneurals 11 (1). Coloration in alcohol. Top of head, snout, supraorbital, sixth and sometimes fifth infraorbitals, and upper half of opercle light-gray to dark-brown. Dorsal portion of body light-gray to dark-brown. Second, third, fourth, and fifth infraorbitals, and opercle silvery in specimens that retained guanine, light- to dark-brown in specimens that lost this pigment due to a long storage in formalin. Dentary, maxillary, gular area, and lower portion of body lightbrown. Lateral portion of body light brown, with a silvery hue in specimens retaining guanine. Humeral blotch present, conspicuous, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of second, 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. Broad, conspicuous longitudinal dark stripe extending at the level of lateral line from supracleithrum to caudal-peduncle and into middle caudal-fin rays to caudal-fin margin. Dark stripe two to two and half scales high. Some relatively small specimens (INHS 60221, CAS (SU) 53787, CAS 64340) with a relatively faint, poorly developed stripe, only developed after at the level of dorsal-fin terminus. Caudal-fin rays with moderately conspicuous, short, elongate, tiny vertical stripes on fin rays. Remaining fins clear, pectoral fins darkened in larger specimens. Coloration in life. Description based on pictures published by Géry (1977: 325, upper picture, as Brycon sp.), Lilyestrom & Taphorn (1983: 55), and pictures from specimens collected at the Río Negro (tributary of Río Casanare), Colombia, by A. Linares. Overall ground color clear, with a silvery hue, top of head, snout and dorsum gray. Longitudinal dark stripe conspicuous, more evident at the level of dorsal fin posteriorly towards caudal peduncle. Pectoral fin dark gray. Dorsal and adipose fins gray. Pelvic, anal, and caudal-fins pinkish. Sexual dimorphism. Anal fin displaying numerous (c. 15 per fin-ray main branch) small hooks on last unbranched and posterior main branch of branched rays 1 – 8 to 10, associated with dense, gelatinous tissue in two specimen (MZUSP 105834, 250.0 – 254.0 mm SL). A single hook per ray segment, a few hooks at the anterior branch of some anal-fin rays. One of these specimens was dissected and proved to be a male. Common names. Venezuela: “ Palambra ” (Lilyestrom & Taphorn, 1983; Taphorn, 1992). Colombia: “ bocón ” (Cala, 1977: 7); “ sardinata ”.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F1CFF974EA4FE8BFC1CFA9F.taxon	distribution	Distribution. Endemic from the Río Orinoco basin in Venezuela and Colombia, where it is widespread, with a clear preference for piedmont areas (Taphorn, 1992) (Fig. 65). Ecological notes. Inhabits mountain, piedmont and high llaneran streams and rivers, preferring clear waters, with strong currents (Taphorn, 1992; Rodríguez-Olarte & Taphorn, 2006). It generally prefers river with an intact gallery forest (Taphorn, 1992). Brycon whitei is most often found in schools (Taphorn, 1992), sometimes mixed with small specimens of Salminus sp. (Rodríguez-Olarte & Taphorn, 2006). The diet of the species was studied by Lilyestrom & Taphorn (1983), who examined stomach contents of 68 specimens and found it constituted mainly (82 %) by vegetal matter (seeds, fruits, leaves), with a smaller amount of animal matter (11 %), constituted by terrestrial insects, small fishes and a lizard. Lasso (2005: 100) examined two stomach contents and found crushed seeds and unidentified vegetal matter. The species is highly fecund (more than 170.000 eggs), breeding during the first month of the wet season (Winemiller, 1989 a; Taphorn, 1992). Brycon whitei undertakes upstream migrations (“ ribazones ”) in the early dry season (October / December) and downstream migrations during the onset of the rainy season (May to June) (Taphorn, 1992). Brycon whitei is highly ranked for both recreational and subsistence fisheries (Lilyestrom & Taphorn, 1983; Taphorn, 1992). The species is reported to reach 450 mm SL and 4 kg (Taphorn, 1992), being consequently one of the largest species in the genus.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F1CFF974EA4FE8BFC1CFA9F.taxon	description	Conservation. Loss of habitat due to deforestation, damming, pollution and water diversion for irrigation projects has caused considerable declines and even local extinctions of populations of Brycon whitei in the piedmont and high llanos area of Venezuela (Lilyestrom & Taphorn, 1983; Taphorn, 1992).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F1CFF974EA4FE8BFC1CFA9F.taxon	discussion	Remarks. Brycon whitei is a distinctive species that has been generally correctly identified since its original description. The only obvious misidentification present in the literature is the record by Saul (1975) of Brycon cf. whitei from the Río Conejo, a tributary of Río Aguarico, Napo, Ecuador. Although the specimens basing that record were not examined during the present study, this record almost certainly refers to Brycon hilarii, a similarlooking species known from the Río Napo basin (see under this species).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F1CFF974EA4FE8BFC1CFA9F.taxon	materials_examined	Material examined. Type material. CAS (SU) 48818 (1, 343.5 mm SL): Colombia, Estado Meta, Río Guaviare [= actually Río Güejar according to maps], vicinity of Los Micos, c. 3 ° 20 ’ N, 73 ° 56 ’ W; T. D. White, J. N. Reynolds & L. Wulff, Feb 1956. Holotype of Brycon whitei Myers & Weitzman. CAS (SU) 48817 (1, 238.0 mm SL): same data as holotype. Paratype of Brycon whitei Myers & Weitzman. Non types. Colombia: CAS (SU) 53787 (1, 151.5 mm SL): Estado Meta, Río Guayabero, Cordillera Macarena, small brook 3 mi. below El Refugio, 2 ° 16 ' N, 73 ° 46 ' W; T. D. White & G. S. Myers, 24 Feb 1960. Venezuela, estado Cojedes: INHS 60211 (1, 192.5 mm SL): Río Pao (Río Portugueza - Río Apure dr.), W La Yeguera, at Hwy. Bridge, 9 ° 32 ’ 25 ’’ N 68 ° 6 ’ 54 ’’ W; L. M. Page et al., 22 Dec 1990. Estado Guárico: MZUSP 106459 (ex MCNG 24231) (2, 143.7 – 157.2 mm SL): caño 8.5 km S of Palenque, río El Burro, 8 ° 47 ’ N, 66 ° 52 ’ W; L. G. Nico et al., 11 Sept 1989. FMNH 85445 (2, 120.7 – 123.2 mm SL): Rio Manapire drainage, 113.1 km south of Chaguaramas, 2.5 km east of highway, tributary of Rio Manapire, 8 ° 20 ’ N, 65 ° 7 ’ W; J. E. Thomerson, D. Hicks, D. Taphorn, T. & D. Greenfield, 5 Jan 1975. Estado Portuguesa: MZUSP 106460 (ex MCNG 90) (2, 151.8 – 177.8 mm SL): Río Las Marias, 800 m above bridge, 4 Km from Guanare, 9 ° 5 ’ 40 ’’ N, 69 ° 30 ’ 20 ’’ W; D. C. Taphorn et al., 26 Feb 1979. CAS 64340 (2, 174.7 – 178.6 mm SL): Río Maria at bridge on Guanare-Acarigua highway, c. 9 ° 5 ’ N, 69 ° 30 ’ W; D. Taphorn et al., 16 Feb 1981. MZUSP 105834 (2, 250.0 – 254.0 mm SL): Río Tucupido, trib. Río Guanare, 15 km SW Guanare, 8 ° 54 ’ 51 ’’ N, 69 ° 45 ’ 40 ’’ W; N. K. Lujan et al., 17 March 2010. Estado Barinas: INHS 61310 (1, 191.0 mm SL): Río Canaguá (Río Apure drainage), 10 km NNW Ciudad Bolivia, Rt. 5 bridge, 8 ° 25 ’ 37 ’’ N, 70 ° 38 ’ 21 ’’ W; L. M. Page et al., 10 Jan 1992. USNM 219726 (4, 264.9 – 306.8 mm SL): Rio Las Palmas (trib. Río Paguey), c. 8 ° 4 ’ N, 70 ° 20 ’ W; P. Bottome & Wallis, 15 Jul 1958.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F18FF784EA4FA0BFE14FD4F.taxon	description	? Brycon stubelli (not Steindachner): Amaral Campos, 1950: 142 (rio Juruá, Amazonas).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F18FF784EA4FA0BFE14FD4F.taxon	description	Brycon cf. falcatus (not Müller & Troschel): Fernández et al., 2006: 55 (photo; upper Río Cataniapo, Rio Orinoco basin, estado Amazonas, Venezuela).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F18FF784EA4FA0BFE14FD4F.taxon	diagnosis	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.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F18FF784EA4FA0BFE14FD4F.taxon	description	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 11 th (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 24 th (1), 25 th (1), or 26 th (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).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F18FF784EA4FA0BFE14FD4F.taxon	distribution	Distribution. Widespread in northern cis-andean South America, in the Rio Amazonas and Rio Orinoco basins (Fig. 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. 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, 1983 a, b; as Brycon cf. melanopterum), an adaptation in fact shared with some other characids (Winemiller, 1989 b). 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).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F18FF784EA4FA0BFE14FD4F.taxon	discussion	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.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F18FF784EA4FA0BFE14FD4F.taxon	description	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. 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. 70 and 75), and thus, we consider the earlier as a synonym of the latter. Steindachner (1879 a: 150) briefly described Brycon longiceps, without mentioning its type-locality. A little later, a proper description of the species was published (Steindachner, 1879 b: 156 – 157). Steindachner (1879 b) 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 (1879 b) 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.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F18FF784EA4FA0BFE14FD4F.taxon	description	Braum (1983 a, 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, 1983 a: fig. 1 a) 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. 85 d), the picture of the species (p. 457) depicts instead a specimen of B. melanopterus.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104F18FF784EA4FA0BFE14FD4F.taxon	materials_examined	Material examined. Type material: INPA 3415 (1, 362.0 mm SL): Brazil, Pará, rio Trombetas, igarapé Caxipacoré, c. 0 ° 34 ’ S, 56 ° 45 ’ W; E. Ferreira & M. Jégu, 20 April 1985. 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 98 (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 8058 (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 date. Holotype of Brycon capito Cope. NMW 62944 (49.1 – 67.7 mm SL): “ Orinoco bei Ciudad Bolivar ” [Venezuela, Estado Bolivar, 8 ° 8 ’ N, 63 ° 34 ’ W]; Schilling, 1879; syntypes of Brycon longiceps Steindachner. FMNH 53353 (1, 169.1 mm SL): Guyana, Mud Creek in Aruka River [c. 8 ° 13 ’ N, 59 ° 44 ’ W]; S. E. Shideler, 1908. Holotype of Brycon siebenthalae Eigenmann. Non types. Brazil, Pará state, rio Amazonas basin: MCZ 21134 (1, 174.7 mm SL); MCZ 21089 (3, 161.4 – 174.0 mm SL): Rio Xingu, Porto de Moz, 1 ° 45 ’ S, 52 ° 14 ’ W; Thayer Expedition, Aug – Sept 1865. BMNH 1926.10. 27.7 (1, 190.4 mm SL): Monte Alegre, Rio Amazonas, 2 ° 0 ’ S, 54 ° 4 ’ W; C. Ternetz, no date. MZUSP 9501 (1, 121.6 mm SL): Monte Alegre, rio Amazonas, 2 ° 0 ’ S, 54 ° 4 ’ W; EPA, 14 Aug 1968. MNRJ 11263 (1, 200.8 mm SL): rio Amazonas, “ Maicura ” (probably rio Maicuru, c. 2 ° 9 ’ S, 54 ° 12 ’ W); L. Travassos & J. de Freitas, Feb 1960. MZUSP 9177 (1, 185.0 mm SL): Santarém, rio Maicá, c. 2 ° 27 ’ S, 54 ° 40 ’ W; EPA, 19 – 27 Oct 1971. CAS 68830 (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 1923. CAS 68884 (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 1924. ZUEC 8033 (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. T. Lima, W. G. R. Crampton, J. S. Ready et al., 22 Nov 2013. MCZ 21099 (2, 139.2 – 170.7 mm SL); MCZ 21098 (1, 155.7 mm SL): Rio Amazonas, Óbidos, 1 ° 52 ’ S, 55 ° 30 ’ W; Thayer Expedition, Nov – Dec 1865. MPEG 11181 (1, 190.6 mm SL): Juruti, Igarapé Guaraná, 2 ° 29 ’ 19 ’’ S, 56 ° 14 ’ 19 ’’ W; A. Hercos & V. Sena, 21 Aug 2006. CAS 66852 (1, 94.3 mm SL): Juruti, 2 ° 9 ’ S, 56 ° 6 ’ W; no collector specified. ANSP 82285 (1, 207.0 mm SL): “ Para ”; J. L. Troemner, no date. Pará state, rio Tapajós basin: MZUSP 8405 (3, 146.6 – 156.6 mm SL): Santarém, igarapé Jacundá, Alter do Chão, c. 2 ° 30 ’ S, 54 ° 58 ’ W; EPA, 23 Dec 1967. MZUSP 57475 (1, 258.1 mm SL): Santarém, Alter do Chão, 2 ° 28 ' 5 '' S 54 ° 55 ' 34 '' W; M. Westneat et al., 31 Oct 1994. MZUSP 60323 (1, 154.5 mm SL): Santarém, rio Tapajós, 2 ° 25 ’ S, 54 ° 45 ’ W; A. Campos, Oct 1944. FMNH 92083 (1, 159.6 mm SL): Santarém, rio Tapajós, 2 ° 25 ’ S, 54 ° 45 ’ W; J. D. Haseman, 12 Dec 1909. MCZ 20780 (1, 117.7 mm SL); MCZ 21094 (1, 176.3 mm SL): Santarém, rio Tapajós, 2 ° 25 ’ S, 54 ° 45 ’ W; D. Bourget, 1865. MCZ 21125 (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 68854 (1, 128.4 mm SL): market at Santarém; C. Ternetz, Aug – Sept 1924. CAS 68837 (1, 243 mm SL); CAS 68915 (3, 115.7 – 121.3 mm SL): rio Tapajós, Santarém, 2 ° 25 ’ S, 54 ° 45 ’ W; C. Ternetz, July 1924. MNHN 1909 - 070 (1, 115.1 mm SL): Santarém, 2 ° 25 ’ S, 54 ° 45 ’ W; C. Jobert, no date. Pará state, rio Trombetas basin: MZUSP 8255 (1, 151.0 mm SL): rio Trombetas, Oriximiná, 1 ° 47 ’ S, 55 ° 52 ’ W; EPA, 16 – 18 Dec 1967. MZUSP 56777 (3, 207.2 – 231.6 mm SL): rio Trombetas, Cuminá, c. 1 ° 31 ’ S, 56 ° 2 ’ W; M. Goulding, Oct – Nov 1983. INPA 16449 (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 1985. INPA 16440 (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 1985. INPA 3416 (1, 317.6 mm SL): rio Mapuera (trib. rio Trombetas), Cachoeira da Égua, c. 1 ° 5 ’ S, 57 ° 20 ’ W; E. Ferreira, 17 May 1985. Amazonas state, rio Solimões / Amazonas basin: MZUSP 5801 (2, 317.2 – 329.7 mm SL): Lago Saracá, Silves, c. 2 ° 53 ’ S, 58 ° 21 ’ W; EPA, 17 – 18 March 1967. MCZ 21124 (1, 176.5 mm SL): Lago Saracá at Silves, c. 2 ° 53 ’ S, 58 ° 21 ’ W; S. V. R. Thayer, Dec 1865. MCZ 21086 (2, 147.0 – 183.2 mm SL): Rio Amazonas, Parintins, 2 ° 38 ’ S, 56 ° 45 ’ W; L. Agassiz, 27 Aug – 2 Sept 1865. MCZ 21091 (1, 160.4 mm SL): Lago José Assú [= Igarapé Açu or rio Andirá, 2 ° 54 ’ S, 57 ° 8 ’ W]; L. Agassiz, 27 – 30 Aug 1865. MZUSP 13430 (1, 323.2 mm SL): Itacoatiara, rio Amazonas, 3 ° 9 ’ S, 58 ° 27 ’ W; N. Smith, 28 Sept 1977. 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 - 1087 (1, 171.5 mm SL): rio Urubu, Salto Lindóia, c. 2 ° 37 ’ S, 59 ° 22 ’ W; M. Jégu, Sept – Oct 1993. MCZ 92873 (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 16450 (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 1977. INPA 4599 (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 1986. 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 16444 (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 13318 (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 1993. 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 307024 (6, 27.9 – 86.2 mm SL): rio Solimões, Ilha de Marchantaria, c. 3 ° 14 ’ S, 59 ° 55 ’ W; P. Bayley, 1977 – 1979. 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 16451 (4, 37.4 – 54.9 mm SL); INPA 16364 (1, 156.7 mm SL): rio Solimões, Lago Janauacá, c. 3 ° 24 ’ S, 60 ° 18 ’ W; Eq. Ictiologia / INPA, Jan 1977. ZUEC 7823 (1,159.1 mm SL): Manaquiri, Lago Janauacá, c. 3 ° 24 ’ S, 60 ° 18 ’ W; V. S. Uieda, 11 – 12 Aug 1979. ZUEC 5345 (1, 89.9 mm SL): Manaquiri, Lago Janauacá, c. 3 ° 24 ’ S, 60 ° 18 ’ W; U. Caramaschi, July 1977. MZUSP 6627 (3, 123.1 – 149.5 mm SL): igarapé tributary of Lago Manacapuru, c. 3 ° 17 ’ S. 60 ° 54 ’ W; EPA, 13 Nov 1967. MZUSP 5903 (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 6306 (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 5990 (1, 70.5 mm SL): rio Purus, at its mouth, c. 3 ° 43 ’ S, 61 ° 27 ’ W; EPA, 1 – 5 May 1967. INPA 17122 (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 2001. INPA 19106 (1, 178.0 mm SL); MCP 29759 (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 1997. MCP 29757 (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 1997. MZUSP 27924 (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 1983. INPA 19108 (1, 139.6 mm SL): rio Solimões, Reserva Mamirauá, Lago Mamirauá; W. G. R. Crampton, Oct 1993. INPA 19118 (1, 39.0 mm SL): rio Solimões, Reserva Mamirauá, Lago Secretaria; W. G. R. Crampton, March 2001. INPA 19107 (2, 119.6 – 133.5 mm SL): rio Solimões, Reserva Mamirauá, lago Curuçá; W. G. R. Crampton, 4 Nov 1997. MCP 29758 (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 1999. INPA 19111 (1, 159.6 mm SL): Tefé, rio Solimões, Lago Capivara, Costa das Capivaras; W. G. R. Crampton, 13 Jan 2001. MCZ 162214 (1, 196.0 mm SL): Tefé, 3 ° 24 ’ S, 64 ° 45 ’ W; L. Agassiz et al., Oct 1865. INPA 19109 (1, 167.1 mm SL): rio Tefé, Ilha do Martelo; W. G. R. Crampton, 14 July 1999. MNHN 1909 - 0289 (1, 153.5 mm SL): Rio Solimões, Tonantins, 2 ° 52 ’ S, 67 ° 46 ’ W; C. Jobert, no date. MZUSP 55821 (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 1993. MZUSP 56153 (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 1993. MZUSP 27334 (2, 236.0 – 237.0 mm SL): Benjamin Constant, Costa do Capacete, rio Solimões, c. 4 ° 20 ’ S, 69 ° 58 ’ W; L. P. Portugal, 13 Nov 1982. MZUSP 27330 (1, 184.0 mm SL): Tabatinga, Lago Caial, rio Solimões, c. 4 ° 20 ’ S, 69 ° 58 ’ W; L. P. Portugal, 8 Oct 1982. MNRJ 11262 (1, 209.5 mm SL): rio Queixito, at its mouth at rio Javari, 4 ° 20 ’ S, 70 ° 12 ’ W; A. Parko, 1942. Amazonas state, rio Uatumã basin: INPA 5685 (1, 258.0 mm SL): rio Uatumã, Balbina, c. 1 ° 55 ’ S, 59 ° 28 ’ W; M. Jégu, Sept 1985. INPA 15421 (2, 286.1 – 292.8 mm SL): rio Uatumã, Balbina, c. 1 ° 55 ’ S, 59 ° 28 ’ W; P. Chaves, July 1985. INPA 16433 (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 5517 (1, 274.7 mm SL): rio Uatumã, igarapé do Arraia; S. Amadio, May 1985. INPA 16428 (1, 287.5 mm SL): igarapé do Arraia, rio Uatumã; Eq. Ictiologia / INPA, 15 April 1983. INPA 16429 (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 1983. INPA 16374 (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 state, rio Negro basin: MCZ 15995 (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 date. MZUSP 9574 (1, 178.3 mm SL): Manaus (fish market); EPA, 17 – 19 Sept 1968. MZUSP 6766 (1, 206.0 mm SL): Manaus, igarapé Tarumãzinho, trib rio Negro, c. 3 ° 3 ’ S, 60 ° 7 ’ W; EPA, 17 – 18 Nov 1967. INPA 15925 (1, 344.0 mm SL): rio Negro, Maruauru (not located); Cidomar, March 2000. MZUSP 61882 (5, 394.5 – 412.5 mm SL): rio Puduari, upper portion, c. 2 ° 8 ’ S, 61 ° 15 ’ W; G. Borges, Nov 1981. 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 2614 (1, 340.9 mm SL): rio Negro, Praia Grande, c. 2 ° 31 ’ S, 69 ° 59 ’ W; Eq. Ictiologia, 2 Jan 1984. UMMZ 217848 (1, 221.5 mm SL): Arquipélago de Anavilhanas, rio Negro; W. F. Fink et al., Jan 1983. UMMZ 217803 (1, 229.4 mm SL): rio Negro, lago “ Chiclaua ” (not located); W. L. Fink, 10 Jan 1983. INPA 9900 (1, 254.7 mm SL): rio Jaú, igarapé Miratuca, c. 1 ° 58 ’ S, 61 ° 30 ’ W; M. Garcia & Oliveira, 29 Oct 1994. 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 1976. INPA 16447 (13, 222.6 – 316.8 mm SL): rio Negro, “ Ponta da Piraíba ” (not located); Eq. Ictiologia / INPA, 15 Aug 1985. MNRJ 6268 (1, 147.4 mm SL): rio Negro, “ Uyipiranga ” (not located); A. Parko, 1941. Amazonas state, rio Madeira basin: MZUSP 7039 (11, 169.8 – 269.7 mm SL): rio Canumã, c. 4 ° 2 ’ S, 59 ° 6 ’ W; EPA, 28 – 29 Nov 1967. MZUSP 59001 (1, 270.8 mm SL): mouth of rio Madeira (purchased at the Itacoatiara fish market); A. M. Zanata, 3 Aug 1996. MCZ 21090 (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 14017 (4, 301.0 – 348.6 mm SL): rio Machado, at its mouth, 8 ° 4 ’ S, 62 ° 53 ’ W; M. Goulding, 18 Sept 1977. UF 100627 (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 16383 (11, 133.0 – 156.8 mm SL): rio Jamari, Samuel dam, diverting channel (cofferdam), c. 8 ° 45 ’ S, 63 ° 27 ’ W; G. M. Santos, 7 – 9 June 1988. INPA 16456 (1, 383.4 mm SL): rio Jamari, 5 km above Samuel dam, c. 9 ° 32 ’ S, 63 ° 8 ’ W; G. M. Santos, 14 July 1985. INPA 16442 (2, 282.8 – 303.2 mm SL): rio Jamari, above Samuel dam, c. 9 ° 32 ’ S, 63 ° 8 ’ W; G. M. Santos, 7 Dec 1984. INPA 16443 (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 96131 (1, 207.0 mm SL): Rio Madeira, “ Cachoele de Theotone ” [= Cachoeira do Teotônio, 8 ° 52 ’ S, 64 ° 3 ’ W]; J. D. Haseman, 30 Oct 1909. CAS 18395 (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 1909. MZUSP 88028 (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 2002. MCP 39586 (1, 188.4 mm SL): Igarapé Taquarás, trib. Rio Mamoré, BR- 425, 9 ° 57 ’ 43 ’’ S, 65 ° 17 ’ 45 ’’ W; A. R. Cardoso, V. Bertaco, F. C. T. Lima & J. F. Pezzi da Silva, 25 Jul 2004. INPA 16371 (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 16375 (2, 192.7 – 207.4 mm SL): Guajará-Mirim, rio Pacaás Novos (trib. rio Guaporé), c. 11 ° 5 ’ S, 65 ° 8 ’ W; G. M. Santos, 16 Nov 1984. INPA 16377 (1, 185.5 mm SL): rio Mamoré, mouth of rio Guaporé, Surpresa, 11 ° 53 ’ S, 65 ° 1 ’ W; G. M. Santos, 19 June 1984. AMNH 39956 (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 1964. Mato Grosso state, rio Madeira basin: MZUSP 105602 (2, 420.0 – 485.0 mm SL): rio Verde, trib. rio Guaporé, Brazil / Bolivia border, 14 ° 7 ’ 36 ’’ S, 60 ° 28 ’ 30 ’’ W; O. A. Cantelmo & L. Barbosa, 13 – 21 Aug 2005. Bolivia, rio Madeira basin: FMNH 107242 (1, 445.0 mm SL): Estado Pando, Río Nareuda (trib. Río Orthon, Río Madre de Dios drainage), c. 11 ° 16 ’ S, 69 ° 3 ’ W; N. A. Menezes et al., 4 Sept 1996. MNHN 1989 – 1430 (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 1984. Peru, Depto. Loreto, Rio Ucayali / Amazonas basin: NRM 23671 (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 1986. MZUSP 15266 (2, 87.2 – 90.7 mm SL): Caño Lupuna, Río Amazonas (not located); M. V. Correa, 10 Jun 1980. INHS 55030 (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 1999. INHS 43857 (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 106469 (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 68880 (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 136578 (1, 164.3 mm SL): Cano del Shansho, near Pebas, c. 3 ° 21 ’ S, 71 ° 48 ’ W; W. G. Scherer, Sept 1936. 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 Tipishca, Río Marañon; H. Ortega et al., 15 Apr 2001. Depto. Ucayali, Río Ucayali basin: MZUSP 26219 (3, 58.9 – 60.8 mm SL); MZUSP 26218 (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 1973. MUSM 41154 (1, 148.8 mm SL): Pucallpa, Río Ucayali, Masisea, 8 ° 35 ’ S, 74 ° 20 ’ W; H. Ortega, 24 Sept 1975. MUSM 108 (2, 133.0 – 134.0 mm SL): Ucayali, Pucallpa, Masisea, Lobococha, 8 ° 35 ’ S, 74 ° 20 ’ W; H. Ortega, 20 Apr 1983. MUSM 315 (2, 130.5 – 132.0 mm SL): Pucallpa, Río Ucayali, Utiquinía, 8 ° 12 ’ S, 74 ° 34 ’ W; H. Ortega, 20 Sept 1985. Colombia, Rio Amazonas basin: UMMZ 191053 (1, 225.5 mm SL): Leticia, Rio Amazonas, 4 ° 13 ’ S, 69 ° 57 ’ W; R. Faber, 17 May 1971. Venezuela, rio Orinoco basin, Estado Delta Amacuro: USNM 236562 (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 1979. Estado Anzoategui: ANSP 166777 (1, 172.0 mm SL): Soledad, Laguna Curita, 8 ° 9 ’ 30 ’’ N, 63 ° 35 ’ 55 ’’ W; M. Rodriguez, 27 Apr 1988. ANSP 166480 (4, 149.5 – 170.9 mm SL): Soledad, Laguna Aguilera, 8 ° 11 ’ 30 ’’ N, 63 ° 26 ’ 45 ’’ W; M. Rodriguez & S. Richardson, 22 Jan 1987. Estado Guárico: MZUSP 54583 (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 1968. INHS 33942 (3, 122.8 – 137.7 mm SL): Pozo Azul (Río San Bartolo), P. N. Aguaro-Guariquito; W. Mejo et al., 4 Jan 1995. INHS 34335 (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 1995. FMNH 85402 (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 1975. LBP 10224 (4, 128.2 – 148.1 mm SL): Cabruta, Río Apure, 7 ° 37 ’ 24 ’’ N, 66 ° 24 ’ 48 ’’ W; C. Oliveira & V. Tagliacollo, 21 Apr 2010. Estado Barinas: FMNH 103954 (1, 154.6 mm SL): Playa Los Chicos in the Rio Suripá, ca. 2.5 hrs. above Hato Las Mercedes, c. 7 ° 47 N, 70 ° 9 ’ W; B. Chernoff et al., 12 Jan 1991. Estado Apure: INHS 27660 (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 1992. INHS 61402 (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 1992. ANSP 165393 (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 1989. ANSP 165548 (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 1989. ANSP 165483 (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 1989. FMNH 69900 (1, 159.3 mm SL); FMNH 117234 (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 1967. Estado Bolívar: ANSP 159722 (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 1985. ANSP 159726 (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 1985. ANSP 159720 (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 1985. ANSP 159727 (1, 117.3 mm SL): Río Caura at Maripa ferry crossing, 7 ° 27 ’ N, 65 ° 12 ’ W; B. Chernoff et al., 19 Nov 1985. ANSP 159728 (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 1985. ANSP 135720 (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 1977. ANSP 135840 (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 1977. CAS uncat. (1, 152.0 mm SL); CAS 156831 (3, 134.6 – 167.9 mm SL): Caño de Quiribana, near Caicara, c. 7 ° 36 ’ N, 66 ° 11 ’ W; C. Ternetz, April – May 1925. CAS 154565 (1, 156.1 mm SL); CAS 154568 (1, 134.9 mm SL): Rio Orinoco, mouth of Cano de Quiribana, Caicara, c. 7 ° 36 ’ N, 66 ° 11 ’ W; C. Ternetz, 5 May 1925. CAS 156758 (1, 130.1 mm SL): creeks into Laguna San Raphael, Caicara, c. 7 ° 36 ’ N, 66 ° 11 ’ W; C. Ternetz, 28 Apr 1925. LBP 3026 (8, 113.7 – 141.8 mm SL): Caicara del Orinoco, Río Orinoco, 7 ° 38 ’ 12 ’’ N, 66 ° 19 ’ 4 ’’ W; C. Oliveira & A. Granado, 2 Oct 2005. ANSP 159740 (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 159725 (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 1985. ANSP 159724 (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 159737 (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 1985. ANSP 159723 (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 1985. ANSP 159739 (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 1985. ANSP 159735 (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 1985. ANSP 135771 (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 1977. 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. 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 85678 (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 159738 (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 1985. FMNH 103953 (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 1992. Guyana: AMNH 14405 (2, 106.5 – 106.9 mm SL); AMNH 14323 (2, 101.3 – 123.9 mm SL): Santa Rosa, Amacuro River, 7 ° 39 ’ N, 58 ° 57 ’ W; A. S. Pinkus, 1938. Trinidad: USNM 235526 (1, 206.3 mm SL): Chatam stream, south west Trinidad, 10 ° 5 ’ N, 61 ° 45 ’ W; M. Atkins, 5 Sept 1980. Uncertain locality: MZUSP 3815 (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 54008 (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 82430 (3, 111.6 – 141.4 mm SL): São Paulo, Campinas, artificial pond near rio Atibaia; A. M. Zanata & M. P. Geraldes, March 2003. MZUSP 86779 (1, 217.5 mm SL): Rio de Janeiro, Campos, rio Paraíba do Sul (fish market); O. T. Oyakawa et al., Sept 2004.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF7FF7D4EA4FD3BFB0CFEA7.taxon	description	Brycon sp.: Santos et al., 1984: 42 (Lower rio Tocantins; brief description, diet).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF7FF7D4EA4FD3BFB0CFEA7.taxon	description	Brycon amazonicus (not Agassiz, in Spix & Agassiz): Mérona et al., 2010: 106, 194 (Lower rio Tocantins, Pará; abundance, impacts of Tucuruí dam). Brycon af. melanopterus (sic) (not Cope): Iglesias-Rios, 2012: 326 (Brazil, Goiás, Serra da Mesa dam: impacts of damming)	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF7FF7D4EA4FD3BFB0CFEA7.taxon	diagnosis	Diagnosis. Brycon gouldingi is distinguished from all its congeners, except B. falcatus, B. melanopterus, B. whitei, B. orbygnianus, B. orthotaenia, B. hilarii, and B. amazonicus 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 gouldingi 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 B. gouldingi is B. amazonicus, from which it can be distinguished by the presence of a distinct crescent- or V-shaped blotch on the caudal peduncle and caudal fin (vs. dark pigment on the caudal peduncle and caudal fin diffuse, never crescent- or V-shaped). See the item “ Comparisons ” on Brycon amazonicus for more information on the diagnosis of both species.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF7FF7D4EA4FD3BFB0CFEA7.taxon	description	Description. Morphometric data are presented in Table 19. One of the largest species in the genus, largest specimen examined reaching 477.9 mm SL. Body moderately slender, moderately high in specimens above 300 mm SL. Largest body height at dorsal-fin level. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, straight to slightly concave from this point to supraoccipital tip, moderately convex from this point to dorsal-fin origin, straight along dorsal-fin base, slightly convex from dorsal-fin terminus 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. Maxilla extending posteriorly to vertical slightly anterior to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Nine (9), 10 (11), 11 (11), 12 (13), or 13 (5) tricuspidate teeth in outer series. Four (11), 5 (30), or 6 (7) trito pentacuspidate teeth in second, inner premaxillary row, plus 3 (2), 4 (40), or 5 (6) tricuspidate teeth between the first and third rows. Two (3 in one specimen) teeth in third premaxillary row, medial teeth largest, pentacuspidate, symphyseal teeth smaller, tri- to pentacuspidate. Maxillary margin approximately straight, except for slight convexity in its distal extremity. Eleven to 21 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 8 (1), 9 (9), 10 (7), 11 (1), or 12 (2) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, penta-, tri- to unicuspidate. Inner (lingual) series consisting of a single symphyseal tooth, considerably smaller than teeth of main series anterior to it, plus row of 15 small, unicuspidate teeth, originating at level of 6 th tooth of main series. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Sixth-six (1), 69 (1), 71 (1), 72 (4), 73 (5), 74 (8), 75 (5), 76 (7), 77 (5), 78 (5), 79 (2), 80 (3), 81 (1), or 82 (1) scales in lateral line series; laterosensory tube ramified, each lateral line scale with two to three branchs in the smallest specimen examined (MZUSP 18196, 118.5 mm SL), gradually becoming more complex during ontogeny, up to the development of ascendent and descendent main branches, and about 6 to 10 secondary branches in specimens larger than 300 mm SL. Horizontal scale rows between dorsal-fin origin and lateral line 13 (6), 14 (24), 15 (14), or 16 (3). Horizontal scale rows between lateral line and pelvic-fin insertion 7 (2), 8 (17), 9 (22), or 10 (6). Circumpeduncular scales 20 (2), 21 (3), 22 (20), 23 (19), 24 (3), or 25 (1). Dorsal-fin rays ii, 9. Dorsal fin origin approximatelly at middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 14 th vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 18 (1), 19 (1), 20 (2), 21 (4), 22 (16), 23 (13), or 24 (10). First anal-fin pterygiophore inserting behind haemal spine of 28 th vertebra. Pectoral-fin rays i, 12 (4), 13 (22), 14 (20), or 15 (2); pelvic-fin rays i, 6 (3), 7 (43), or 8 (2); main caudal-fin rays 10 / 9. Last unbranched and anterior 2 – 3 anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin presenting small hooks on posterior branch of branched rays in one specimen (MZUSP 70420). Sheath of scales composed of 4 to 5 horizontal series, covering basal third of analfin rays. Caudal fin forked, lobes slightly emarginated. Four branchiostegal rays. First gill arch with 15 (3), 16 (3) or 17 (3) lower gill rakers, 16 (2), 17 (5), or 18 (2) upper gill rakers and 1 at angle. Vertebrae 47 (1). Supraneurals 10 (1). Coloration in alcohol. Overall color pattern of specimens still retaining guanin light gray dorsally, with silvery hue. Top of head and snout dark gray; infraorbitals and opercle silvery. Maxilla, gular area, and dentary light brown. Dorsolateral body surfaces silvery to coppery, dark brown dorsally, becoming gradually clear ventrally. Humeral blotch present, oval, horizontally elongated, formed by subjacent pigment, lying above lateral line, at level of second to fourth scales. Longitudinal stripes extending all along the trunk, wavy, formed by dark pigment concentrated on upper and lower scale margins in dorsolateral surfaces of body. Relatively faint, rounded dark area on caudal peduncle. Caudal pigmentation contiguous with caudal peduncle blotch, occupying most of caudal fin except for distal margin and two dorsalmost and two ventralmost principal rays. Contour of blotch formed by dark pigmentantion on caudal peduncle and caudal fin crescent or V-shaped. Moderate to heavy darkly pigment on pectoral fins, pigmentation mainly concentrated on outermost seven to eight rays. Pelvic fins pale to intensely dark. Intense dark pigmentation on anal-fin interradial membranes, mainly on posteriormost rays. Dorsal fin pale, with some dark pigmentation on interradial membranes. Adipose fin dark gray. Specimens which lost guanine as a result of long storage in formalin with overall color pattern brownish, without silvery hues on dorsolateral surfaces of body, infraorbitals, or opercle, and with longitudinal stripes more evident. Color in life. Description based on slides taken by Carlos Figueiredo from two individuals collected in the rio Tocantizinho basin, upper rio Tocantins, in the region of the present Serra da Mesa dam, and from specimens collected by J. B. Nunes in the rio Araguaia system. Dorsum bluish-gray; top of head and snout dark gray; sides of body, opercle and infraorbitals pale, with metallic hue; humeral blotch and wavy dark stripes on sides of body clearly visible; pectoral fin darkened; crescent-shaped black blotch on caudal peduncle and caudal fin conspicuous. Dorsal and adipose fins reddish, pelvic and anal fins yellowish, with a tenuous, dark basal stripe. Common names. Matrinchã, piabanha (Santos et al., 1984: 42).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF7FF7D4EA4FD3BFB0CFEA7.taxon	distribution	Distribution. Endemic from the rio Tocantins basin, where it is widespread in the main river systems: the rio Tocantins itself, rio Araguaia, rio das Mortes, rio Paranã, and rio Itacaiúnas, Brazil (Fig. 74). In the last few years, Brycon gouldingi has been stocked outside its native range, and escaped specimens were reported from the rio São Francisco in Pernambuco (J. M. Barbosa, pers. comm ..). Mauro C. Lopes (pers. comm ..) photographed a juvenile of the species fished at the rio Xingu in the Parque Indígena do Xingu, Mato Grosso, in 2008. Since extensive collecting activities just above the park, in the rio Culuene, revealed only Brycon falcatus (pers. obs.), it is suspected that this record might be the result of released specimens collected from the neighbouring rio Araguaia basin. At least one rancher at the upper rio Xingu basin was known to stock fishes from the rio Araguaia system. These fishes were released into the rio Suiá-Missu (a tributary of the upper rio Xingu) when the dam where they were stocked burst after major rains (Lício Moraes, pers. comm.). There are still no evidence of established populations outside the natural area of distribution. Ecological notes. Diet mentioned as being composed of fruits and insects (Santos et al., 1984: 42). Examination of the stomach contents of one specimen (MZUSP 18069, 161.3 mm SL), revealed mainly pieces of fruits and crushed seeds, but also a small cockroach and an ant. Albrecht et al. (2009), in a detailed analysis of the diet of the species at Serra da Mesa dam, found, in order of importance, seeds and fruits, terrestrial invertebrates, and fishes as the most important dietary itens for the species. James B. Nunes (pers. comm.) reported that the species undertakes massive migrations in the area of Ilha do Bananal, middle rio Araguaia basin. Brycon gouldingi leave the floodplains at the end of the rainy season in mixed schools with Prochilodus nigricans and Leporinus trifasciatus. Breeding schools are observed during the rainy season, between December and March. Conservation. The rio Tocantins basin is progressively being dammed, with five major hydroelectric dams already in operation (Tucuruí, Estreito, Lajeado, Peixe / Angical, and Serra da Mesa) and a few more being planned. It is expected that, as a presumable long-range total spawner, Brycon gouldingi populations will be severely impacted by these dams. Prospects on the rio Araguaia are a litlle better, since not much hydroelectric development is expected in that river basin due to its general low gradient. Overall, the conservation status of Brycon gouldingi give some causes to concern as long as developmental plans for the Araguaia / Tocantins basin by the Brazilian government remain unabated.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF7FF7D4EA4FD3BFB0CFEA7.taxon	materials_examined	Material examined. Type material. Holotype: MZUSP 61400 (1, 374.7 mm SL): Brazil; Pará; Parauapebas, Igarapé da Boa Vista, tributary of rio Itacaiúnas, Serra dos Carajás c. 5 ° 52 ’ S, 50 ° 29 ’ W; M. Goulding, Nov 1983. Paratypes: Brazil, Pará: MZUSP 56808 (3, 214.8 – 237.2 mm SL); MZUSP 56948 (1, 262.3 mm SL): Parauapebas, Serra dos Carajás, rio Itacaiúnas; Caldeirão, c. 5 ° 52 ’ S, 50 ° 29 ’ W; M. Goulding, April 1983. MZUSP 56949 (3, 313.4 – 365.9 mm SL): Parauapebas, igarapé do Cinzento, rio Itacaiúnas, c. 5 ° 51 ’ S, 50 ° 32 ’ W; M. Goulding, Nov 1983. MZUSP 58044 (1, 224.4 mm SL): Parauapebas, igarapé Salobo, tributary of rio Itacaiúnas, 5 ° 46 ’ 50 ’’ S 50 ° 32 ’ 48 ’’ W; P. S. Pompeu, July 1997. MZUSP 57033 (28, 199.7 – 298.0 mm SL); MCP 34184 (1, 223.5 mm SL); FMNH 112957 (1, 218.0 mm SL); USNM 374028 (1, 220.7 mm SL): Parauapebas, rio Itacaiúnas, Caldeirão, c. 5 ° 52 ’ S, 50 ° 29 ’ W; M. Goulding, April / May 1983.; MZUSP 61399 (1, 353.9 mm SL): Parauapebas, Igarapé Águas Claras, Serra dos Carajás, c. 5 ° 52 ’ S, 50 ° 29 ’ W; M. Goulding, Nov 1983. MZUSP 31449 (1, 345.1 mm SL): Parauapebas, rio Itacaiúnas, Serra dos Carajás, c. 5 ° 52 ’ S, 50 ° 29 ’ W; M. Goulding, Nov 1983. MZUSP 18100 (1, 161.5 mm SL): marginal lake from igarapé Espírito Santo, between Baião and Tucuruí, rio Tocantins; EPA, 11 Sept 1970. MZUSP 18090 (1, 231.2 mm SL): Baião, rio Tocantins, c. 2 ° 48 ’ S, 49 ° 41 ’ W; EPA, 9 – 10 Sept 1970. MZUSP 18078 (2, 152.5 – 186.5 mm SL): igarapé do Limão, near Baião, rio Tocantins, c. 2 ° 48 ’ S, 49 ° 41 ’ W; EPA, 9 Sept 1970. MZUSP 18050 (1, 192.0 mm SL): Paraná Samuuma, mouth of Tocantins, c. 1 ° 56 ’ S, 49 ° 12 ’ W; EPA, 4 Sept 1970. MZUSP 18196 (1, 118.5 mm SL): igarapé dos 5, km 5 from the road Tucuruí-Mato Grosso, c. 3 ° 42 ’ S, 49 ° 27 ’ W; EPA, 22 Sept 1970. MZUSP 18069 (2, 1 cs, 161.3 – 198.9 mm SL): Mocajuba, igarapé Oxipucu, rio Tocantins, 2 ° 34 ’ S, 49 ° 31 ’ W; EPA, 8 Sept 1970. INPA 16370 (1, 226.2 mm SL): rio Tocantins, Tucuruí fish market, c. 3 ° 42 ’ S, 49 ° 27 ’ W; Eq. Ictiologia / INPA, 27 June 1982. INPA 16367 (1, 121.0 mm SL): rio Tocantins, igarapé Vermelho, 0.5 hour above its mouth, Itupiranga, c. 5 ° 09 ’ S, 49 ° 20 ’ W; Eq. Ictiologia / INPA, 2 July 1982. INPA 16393 (1, 107.7 mm SL): Acarí Pucú, rio Tocantins, 2 ° 42 ’ S, 49 ° 43 ’ W; Eq. Ictiologia / INPA, 27 March 1982. INPA 16379 (1, 210.7 mm SL): rio Tocantins, Tucuruí, c. 3 ° 42 ’ S, 49 ° 27 ’ W; Eq. Ictiologia / INPA, 26 Oct 1980. INPA 16376 (3, 158.1 – 184.4 mm SL): rio Tocantins, Itupiranga, c. 5 ° 09 ’ S, 49 ° 20 ’ W; Eq. Ictiologia / INPA, 6 July 1980. INPA 16247 (1, 289.0 mm SL): rio Tocantins, Itupiranga, Ressaca dos Macacos, c. 5 ° 09 ’ S, 49 ° 20 ’ W; Eq. Ictiologia / INPA, 3 July 1982. Non types. Brazil, Tocantins: UNT 2197 (1, 477.9 mm SL): Peixe, rio Santa Tereza, c. 11 ° 47 ’ S, 48 ° 38 ’ W; NEAMB, 12 March 2001. UNT 3442 (2, 92.3 – 116.2 mm SL); UNT 3440 (1, 136.8 mm SL); UNT 3517 (1, 244.0 mm SL): Porto Nacional, rio Tocantins, c. 10 ° 42 ’ S, 48 ° 25 ’ W; NEAMB, Feb – April 2002. INPA 20117 (1, 254.8 mm SL): Caseara, rio Araguaia, Lago Volta Grande, P. E. Cantão, 9 o 47 ’ 19 ” S, 50 o 9 ’ 53 ” W; J. Zuanon et al., 19 May 2000. INPA 20490 (1, 359.0 mm SL): Caseara, lago Paredão, P. E. Cantão, 9 o 22 ’ 40 ” S, 49 o 58 ’ 32 ” W; J. Zuanon et al., 19 Feb 2000. Goiás: MZUSP 44572 (1, 382.2 mm SL): Monte Alegre de Goiás, rio Paranã, above mouth of rio Bezerra, c. 13 ° 16 ’ S, 47 ° 31 ’ W; J. C. Oliveira & W. J. E. M. Costa, 10 – 13 Jan 1989. MZUSP 70423 (1, 283.8 mm SL): rio Tocantins, below Serra da Mesa dam, 13 ° 49 ' 55 ’’ S, 48 ° 17 ' 44 ’’ W; D. F. Moraes et al., 5 Feb 1997. MZUSP 70422 (1, 278.2 mm SL): Serra da Mesa dam, rio Tocantizinho arm, 13 ° 57 ' S, 48 ° 15 ' W; D. F. Moraes and D. A. Halboth, 6 June 1997. MZUSP 70420 (1, 328.1 mm SL): Serra da Mesa dam, rio Palmeirinha arm, 14 ° 3 ' 57 ' ’ S, 48 ° 29 ' 37 ' ’ W; D. F. Moraes and D. A. Halboth, 16 Feb 1995. MZUSP 70421 (1, 251.2 mm SL): Serra da Mesa dam, above rio Castelo Grande, 14 ° 8 ' S, 48 ° 44 ' W; D. F. Moraes, 10 June 1997. MZUSP 70752 (1, 346.2 mm SL): Serra da Mesa dam, rio Tocantizinho arm, 14 ° 00 ’ S, 48 ° 11 ’ W; D. F. Moraes and D. A. Halboth, 5 Feb 1998. MCP 17280 (1, 452, 7 mm SL): Luís Alves, marginal lagoons of the rio Araguaia, 13 ° 14 ' S, 50 ° 35 ' W; F. L. T. Garro, 21 April 1994. MCZ 21126 (1, 355.0 mm SL): “ Goyaz ” (exact locality unknown; see Lima, 2004: 285); “ Honório ” [A. H. Ferreira], 1867. Mato Grosso: MZUSP 48114 (1, 365.4 mm SL): Barra do Garças, Córrego da Ponte Queimada, c. 15 ° 53 ’ S, 52 ° 15 ’ W; W. P. Margarido, 1 Sept 1993. MZUSP 60411 (1, 278.3 mm SL): Barra do Garças, rio Araguaia, Ouro Fino, 30 km below Barra do Garças, 15 ° 52´03 ´´ S, 51 ° 58´17 ´´ W; W. Barrella et al., 6 – 7 Oct 1997.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF2FF754EA4FE53FED6F992.taxon	materials_examined	Brycon hilarii (not Valenciennes, in Cuvier & Valeciennes): La Monte, 1935: 7 (“ Jurua ”; material collected by B. Krukoff, examined). Brycon melampterum (sic): Eigenmann & Allen, 1942: 253 – 254 (Peru: Yarinococha; Orellana, Rio Ucayali; Rio Pachitea; Pto. Bermudez, Rio Pichis; mouth of Río Pacaya).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF2FF754EA4FE53FED6F992.taxon	description	Brycon cephalus (not Günther): Galvis et al., 2006: 457 (photo).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF2FF754EA4FE53FED6F992.taxon	diagnosis	Diagnosis. The presence of an oblique, solid dark stripe extending from immediately behind pelvic-fin basis, through anal-fin basis, lower surface of caudal peduncle, middle area of caudal peduncle, to the upper caudal-fin lobe, diagnoses Brycon melanopterus from all remaining Brycon species, with the exception of B. falcatus and B. amazonicus. Brycon melanopterus can be diagnosed from the often sympatric B. amazonicus by possessing dark, straight longitudinal stripes formed by pigmentation concentrated on the mid-distal portion of scales (vs. dark, wavy longitudinal stripes formed by dark pigment concentrated on upper and lower scale margins), oblique dark stripe continuous and solid dark (vs. oblique stripe, when present, blurred and not continuous), and paired fins clear (vs. paired fins black). Brycon melanopterus is more similar to B. falcatus, and an unequivocal diagnosis between both species is not possible due to the great polymorphism observed in the latter species. However, typically, Brycon falcatus does not present an oblique dark solid stripe, presenting instead a V or crescent-shaped blotch on caudal fin (vs. black pigmentation restricted to the upper caudal-fin lobe in B. melanopterus). Additionaly, Brycon melanopterus is typically a more elongated fish, with higher vertebrae counts. See item “ Comparisons ”, below, for a more detailed comparison with Brycon falcatus.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF2FF754EA4FE53FED6F992.taxon	description	Description. Morphometric data are presented in Table 20. Middle-sized species, largest examined specimen 289.1 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, snout pointed, 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 (7), 9 (12), 10 (31), 11 (14), 12 (13), or 13 (7) relatively small tricuspidate teeth in outer series. Four (26), 5 (48), 6 (15), or 7 (2) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 3 (21), 4 (61), or 5 (9) 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, both pentacuspidate. Maxillary margins approximately parallel, straight in profile. Fifteen to 26 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 8 (3), 9 (8), 10 (17), 11 (8), 12 (8), 13 (4), or 14 (1) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, pentacuspidate, 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 21 – 24 small, aciculated, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of fifth to sixth main series dentary teeth. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Fifty-four (1), 56 (2), 57 (1), 60 (1), 61 (8), 62 (8), 63 (13), 64 (9), 65 (19), 66 (14), 67 (7), 68 (5), or 70 (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 (> 270 mm SL) specimens. Horizontal scale rows between dorsal-fin origin and lateral line 11 (2), 12 (20), 13 (56), 14 (10), or 15 (2). Horizontal scale rows between lateral line and pelvic-fin 5 (8), 6 (46), 7 (32), or 8 (4). Circumpeduncular scales 18 (1), 19 (12), 20 (20), 21 (31), 22 (21), or 23 (5). Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 12 th (1) vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in cs specimens), 19 (3), 20 (2), 21 (11), 22 (29), 23 (27), 24 (13), or 25 (2). First anal-fin pterygiophore inserting behind haemal spine of 24 th (1) 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 19 – 23 rectangular scales. Pectoral-fin rays i, 11 (1), 12 (28), 13 (51), or 14 (11). Pelvic-fin rays typically i, 7 (87), rarely i, 5 (10), or i, 8 (3). Main caudal-fin rays 10 / 9. Caudal fin forked. 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 11 (5), 12 (6), 13 (12), 14 (4), or 15 (5) lower, 1 at angle, and 9 (2), 11 (8), 12 (15), 13 (6), or 14 (2) upper gill rakers. Vertebrae 45 (2). Supraneurals 9 (1). Coloration in alcohol. Top of head, snout, supraorbital, and sixth infraorbital light-grey to light-brown. Dorsal portion of body light-grey 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 silvery in specimens that retained guanine, light brown in specimens that lost this pigment due to a long storage in formalin. Humeral blotch present, slightly to moderately conspicuous, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of second, extending longitudinally to posterior margin of fifth to sixth lateral line scales, and vertically one and half scales high. Scales on lateral portions of body with dark pigment concentrated on their central portion, forming dark, straight longitudinal stripes, more conspicuous dorsally. Series of irregular, narrow vertical stripes present in small (up to 100 mm SL) specimens. Dark, solid, oblique dark stripe extending in most specimens from immediately behind pelvic-fin basis, through anal-fin basis, lower surface of caudal peduncle, middle area of caudal peduncle, to the upper caudal-fin lobe. Some specimens (most of them obviously faded) with dark stripe starting only at anal-fin origin. Lower caudal-fin rays, dorsal and anal-fins clear, with a variable amount of interradial dark pigmentantion. Adipose-fin light- to dark-grey. Color in life. Based on pictures of specimens collected in tributaries of the Rïo Ucayali at Zona Reservada Sierra del Divisor, Ucayali, Río Pacaya, Reserva Nacional Pacaya Samíria, Loreto, Peru, and Río Mesay, Río Caquetá basin, Colombia. Snout and top of head grey. Overall gound color clear, with a silvery hue, coppery on dorsum. Oblique dark stripe extending from pelvic-fin basis to upper lobe of caudal fin very conspicuous. Adipose fin and middle caudal-fin rays with some yellowish pigmentation, remaining fins translucent. Common names. Brazil: “ matrinchão ”, rio Madeira basin, Rondônia; “ jatuarana ”, Manaus and Tefé regions, Amazonas (Goulding, 1980; Borges, 1986; Géry & Mahnert, 1992; Crampton, 1999; Santos et al., 2006). Both names are also applied, but with an inverse usage, to Brycon amazonicus on those same regions (see under “ Common names ” of B. amazonicus). Tefé region: “ gogota ” (Crampton, 1999: 15; Barthem, 1999: 82). Ecuador: “ sábalo ” (Barriga, 1991: 14). Peru: “ sábalo cola negra ” (Ortega & Vari, 1986: 7).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF2FF754EA4FE53FED6F992.taxon	distribution	Distribution. Widespread in western Amazon, at the main channel of the Rio Amazonas / Solimões upstream to Río Ucayali in Peru, Río Napo in Ecuador, and Río Caquetá in Colombia, rio Madeira system upstream to the rio Madre de Dios in Peru, and lower rio Negro and rio Branco systems in Brazil (Fig. 83). Additionally, Géry & Mahnert (1992: 798, fig. 5) portrays a Brycon melanopterus specimen collected in the rio Guaporé at Vila Bela (da Santíssima Trindade), in the upper rio Madeira basin at Mato Grosso state, Brazil. Comparisons. As noticed in the Diagnosis, an unequivocal diagnosis between Brycon melanopterus and B. falcatus is not possible due to the great range of pigmentary and morphological variation observed in the latter species. While typically Brycon falcatus does not possess a solid oblique dark bar extending from pelvic-fin basis to the upper lobe of caudal peduncle as B. melanopterus, presenting instead a black stripe on anal-fin basis not continuous with a V-shaped blotch on caudal-fin and caudal peduncle, some populations of Brycon falcatus possess a color pattern similar or almost identical to B. melanopterus. Brycon melanopterus and B. falcatus are however largely alopatric, occurring sympatrically only at the western Amazon (where B. falcatus is very local) and at the lower rio Negro and rio Branco basins. Populations of Brycon falcatus from western Amazon in Brazil and Peru, sympatric and in some cases collected syntopically with B. melanopterus (INHS 106468, INHS 106470, MZUSP 17530, MZUSP 99213), are very similar to the latter species since they have a reduced amount or lack entirely dark pigmentation on the lower caudal-fin lobe. However, unlike Brycon melanopterus, these specimens possess what could be called an interrupted oblique dark stripe, since the blotch in the caudal peduncle and caudal fin is not continuous with the dark stripe at the basis of anal fin (compare Figs. 5 I and L). Also, although there is a great overlap in morphometric data, Brycon melanopterus is generally a more elongated fish (28.6 – 39.7, mean 32.5 % of SL, vs. 28.8 – 42.8, mean 36.4 in B. falcatus), with a shallower head (73.8 – 91.3, mean 82.3 % of HL, vs. 76.7 – 1.14, mean 90.6 in B. falcatus) than B. falcatus. The fact that Brycon melanopterus is more elongated than B. falcatus is also reflected in its higher vertebrae counts when compared with the latter species (45, n = 2, vs. 39 – 44, modally 41, n = 17, respectively). Populations of Brycon falcatus inhabiting the upper rio Negro basin in Brazil and the upper rio Orinoco basin in Venezuela possess a color pattern that is virtually undistinguishable from the one presented by B. melanopterus, i. e., the oblique bar is continuous from pelvic-fin basis / anal-fin origin to upper lobe of caudal fin. These specimens (MZUSP 91493, MZUSP 91576, FMNH 104026, ANSP 161210, AMNH 93064, AMNH 93065) are here assigned to Brycon falcatus due to the fact that in overall body shape and vertebrae counts they agree more with the latter species than with B. melanopterus. Brycon melanopterus is apparently absent from the upper rio Negro above the São Gabriel da Cachoeira rapids (0 ° 8 ’ S, 67 ° 5 ’ W), and specimens of the species collected in the lower rio Negro and at the rio Branco are morphologicaly undistinguishable from specimens from the Rio Amazonas / Solimões / Ucayali and rio Madeira basins. Sympatric populations of Brycon falcatus in the middle and lower rio Negro basin can be readily distinguished from B. melanopterus by possessing dark pigmentation on the lower caudal-fin lobe (though considerably less intense than the dark pigmentation in the upper caudal-fin lobe), and, as all remaining Brycon falcatus populations, by not possessing an oblique dark stripe, i. e., the caudal peduncle / caudal fin blotch is not continuous with the dark stripe at the basis of anal fin.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF2FF754EA4FE53FED6F992.taxon	description	Ecological notes. Goulding (1980) and Borges (1986) presented information on the ecology of Brycon melanopterus, from respectively the rio Machado (rio Madeira basin, Rondônia state, Brazil) and rio Negro (Amazonas state, Brazil). Goulding (1980: 73 – 75) reported that the species is less abundant and does not grow as large as the sympatric Brycon amazonicus (his Brycon sp.). Dietary itens found in five stomachs included crushed seeds of Tabebuia barbata (Bignoniaceae), Hevea brasiliensis (Euphorbiaceae), and Mabea sp. (Euphorbiaceae), as well as terrestrial arthropods and a fish. Since the fishermen does not report schools of Brycon melanopterus moving in the rio Machado and rio Madeira, Goulding (1980) infered that the species does not undertake longrange, massive reproductive migrations. Borges (1986) reported fruits and seeds as the dominant dietary item in specimens of Brycon melanopterus (his Brycon cf. cephalus) collected in the rio Negro, followed by arthropods, leaves, flowers, mammals and faeces. Mérona & Rankin-de-Mérona (2004: 77) noticed a similar dietary preference for specimens of Brycon melanopterus collected in a floodplain lake in the Amazon river. Among the fruits and / or seeds ingested in the rio Negro by the species, the most abundant were those belonging to the family Euphorbiaceae (notably Alchornea schomburgkiana), but also Annonaceae (mainly Pseudoxandra polypheba), Lauraceae (Ocotea sp.), and Leguminosae (Centrosoma sp. and Swartzia sp.) (Borges, 1986). Correa & Winemiller (2014) reported that the species switched from fruits and seeds to terrestrial invertebrates as its main dietary item from rainy season to dry season in a site at the lower Río Caquetá basin (= Rio Japurá in Brazil) in Colombia.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF2FF754EA4FE53FED6F992.taxon	discussion	Remarks. Cope (1871: 262 – 263) described Megalobrycon melanopterum based on two specimens collected by J. Hauxwell at the “ Ambyiacu ”, in his words “ an inconsiderable river, which empties into the Amazon near to Pebas, in Eastern Ecuador, some distance east of the Napo ”. Currently, the Río Ampyiacu (not “ Ambyiacu ”) lies in the Departamento Loreto in Peru. Fowler (1907: 447) redescribed the species, based on the type specimens and additional specimens collected by J. Orton (ANSP 21972). He cited the two “ co-types ” (= syntypes), and designated the largest specimen (ANSP 8035) as the “ type ” of the species, which can be accepted as a designation of a lectotype (ICZN, 1999, articles 74.3 and 74.5). The identification of the specimens ANSP 8035 and ANSP 8036 as being, respectively, the holotype and paratype of Megalobrycon melanopterum, made by Böhlke (1984: 51) and subsequently followed by some other authors (Eschmeyer, 1998: 1055; Lima, 2003: 177) is thus incorrect, and these specimens should be henceforward referred as the lectotype and paralectotype of the species. The good illustration of the species present in the original description (Cope, 1871; pl. 13, fig. 1) depicts quite well the oblique, continuous dark stripe extending from anal-fin basis to the upper lobe of caudal fin typical for the species. Possibly due to this fact, and relatively unusually in the history of taxonomy of Brycon, the name B. melanopterus was generally correctly employed in the literature (e. g., Eigenmann & Allen, 1942; Knöppel, 1970; Goulding, 1980; Géry & Mahnert, 1984, 1992). However, a few misidentifications have occurred in the literature, mainly involving Brycon amazonicus, a largely sympatric species which occasionaly also displays an oblique dark stripe similar to B. melanopterus (see item “ Variation ” and “ Remarks ” from B. amazonicus). For example, the examination of the lot identified by Amaral Campos (1950: 142 – 143) as Brycon melanopterum (MZUSP 3574) revealed that it was actually composed by both B. melanopterus and B. amazonicus. Also, Howes (1982: 20) incorrectly supposed that the specimen identified by Goulding (1980: 74, fig. 4.7) as Brycon melanopterus was a misidentification of B. amazonicus (his B. cephalus), when in fact Goulding (1980) has correctly identified the species.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FF2FF754EA4FE53FED6F992.taxon	materials_examined	Material examined. Type material: ANSP 8035 (1, 142.6 mm SL): “ Ecuador, Ambyiacu River ” [= Peru, Depto. Loreto, Río Ampyiacu, trib. Río Amazonas, c. 3 ° 21 ’ S, 71 ° 47 ’ W]; J. Hauxwell, no date. Lectotype of Megalobrycon melanopterum Cope, designated by Fowler (1907: 447). Non types. Brazil, Amazonas state, rio Amazonas / Solimões basin: MCZ 21088 (1, 152.4 mm SL): Lago Máximo, near Parintins, 2 ° 40 ’ S, 56 ° 45 ’ W; L. Agassiz et al., 27 – 30 Aug 1865. MZUSP 13429 (1, 280.6 mm SL): Itacoatiara, rio Amazonas, c. 3 ° 9 ’ S, 58 ° 27 ’ W; N. Smith, 29 Sept 1977. MNHN 1996 - 1087 (1, 171.5 mm SL): Rio Urubu, cachoeira Lindóia, c. 2 ° 37 ’ S, 59 ° 22 ’ W; M. Jégu, 23 Sept 1993. MZUSP 6090 (1, 181.2 mm SL): lago Puraquequara, rio Puraquequara mouth, 3 ° 2 ’ S, 59 ° 47 ’ W; EPA, 17 – 19 Apr 1967. INPA 16366 (1, 167.5 mm SL): rio Amazonas, Paraná do Rei, Ilha do Careiro, c. 3 ° 6 ’ S, 59 ° 49 ’ W; C. Cox Fernandes, 17 Sept 1986. INPA 16372 (3, 86.1 – 185.1 mm SL); INPA 16385 (2, 119.8 – 127.2 mm SL): rio Amazonas, Ilha do Careiro, c. 3 ° 6 ’ S, 59 ° 49 ’ W; – Eq. Ictiologia / INPA, June – Oct 1986. MCZ 21087 (1, 127.8 mm SL): Paraná do Janauari, 3 ° 12 ’ S, 60 ° 5 ’ W; L. Agassiz, 27 – 29 Oct 1865. USNM 306848 (5, 70.2 – 101.8); USNM 309178 (1, 124.7 mm SL): Lago Janauari, c. 3 ° 12 ’ S, 60 ° 5 ’ W; P. Bayley, June – July 1977. INPA 13298 (3, 30.3 – 52.2 mm SL); INPA 13304 (3, 47.5 – 65.4 mm SL); INPA 16353 (2, 61.8 – 67.7 mm SL; INPA 13322 (8, 47.3 – 66.0 mm SL): rio Solimões, Ilha da Marchantaria, c. 3 ° 14 ’ S, 59 ° 55 ’ W; P. Petry & R. Sotero, Jan 1993. INPA 16347 (5, 89.0 – 97.1 mm SL); INPA 16453 (2, 44.6 – 58.2 mm SL): rio Solimões, Ilha da Marchantaria, c. 3 ° 14 ’ S, 59 ° 55 ’ W; Eq. Ictiologia / INPA, 1976 – 1981. USNM 307013 (1, 73.1 mm SL); USNM 307012 (1, 73.1 mm SL); USNM 307011 (2, 68.9 – 75.9 mm SL); USNM 307005 (3, 57.4 – 74.8 mm SL); USNM 229140 (2, 67.8 – 75.7 mm SL): Ilha da Marchantaria, c. 3 ° 14 ’ S, 59 ° 55 ’ W; P. Bayley, Jan – Apr 1978. MZUSP 18693 (1, 129.2 mm SL); MZUSP 18699 (1, 46.7 mm SL): Lago Janauacá and surroundings, rio Solimões, c. 3 ° 24 ’ S, 60 ° 17 ’ W; Alpha Helix Exp., Sept 1976 — Jan 1977. MZUSP 75566 (3, 93.3 – 106.8 mm SL); USNM 307073 (3, 99.8 – 121.4 mm SL); USNM 229075 (1, 115.2 mm SL): Lago Murumuru, into Lago Janauacá, c. 3 ° 24 ’ S, 60 ° 17 ’ W; P. Bayley, 28 Sept 1977. MCP 27932 (1, 70.5 mm SL): Lago Murumuru, into Lago Janauacá, c. 3 ° 24 ’ S, 60 ° 17 ’ W; P. Bayley, 31 Aug 1977. MZUSP 6626 (12, 110.3 – 132.7 mm SL): igarapé trib. lago Manacapuru, c. 3 ° 19 ’ S, 60 ° 50 ’ W; EPA, 13 Nov 1967. MZUSP 5880 (1, 241.6 mm SL): Lago Manacapuru, c. 3 ° 19 ’ S, 60 ° 50 ’ W; EPA, 26 – 27 March 1967. MCZ 21095 (1, 153.2 mm SL): Lago Grande de Manacapuru, 3 ° 6 ’ S, 61 ° 30 ’ W; W. James, Nov – Dec 1865. MCZ 21085 (2, 122.1 – 213.0 mm SL): Rio Solimões at Tefé, 3 ° 24 ’ S, 64 ° 45 ’ W; L. Agassiz et al., Oct 1865. MCP 29772 (1, 123.8 mm SL): Tefé, mouth of igarapé Açu, 3 ° 24 ’ 52 ’’ S, 64 ° 48 ’ 7 ’’ W; W. G. R. Crampton, 21 Jan 1999. MCP 29773 (1, 118.9 mm SL): Alvarães, Lago Mamirauá, comunidade Boca do Lago Mamirauá, 3 ° 6 ’ 37 ’’ S, 64 ° 47 ’ 49 ’’ W; W. G. R. Crampton, 1 Nov 1997. MCP 29974 (2, 142.8 – 148.5 mm SL): Alvarães, between Lago Secretário and Lago Mamirauá channel, 3 ° 6 ’ 58 ’’ S, 64 ° 47 ’ 49 ’’ W; W. G. R. Crampton, Oct 1993. INPA 19117 (1, 119.3 mm SL); MCP 29975 (1, 136.2 mm SL): Lago Amanã, mouth of igarapé Baré, 2 ° 28 ’ S, 64 ° 43 ’ W; W. G. R. Crampton, 18 Dec 1997. MZUSP 17616 (2, 221.0 – 235.0 mm SL): igarapé n ° 1, Fonte Boa, 2 ° 30 ’ S, 66 ° 4 ’ W; EPA, 25 Oct 1968. MZUSP 56498 (3, 146.9 – 177.0 mm SL): igarapé da Cachoeira, Cuiauá, rio Içá, left margin, c. 3 ° 0 ’ S, 69 ° 2 ’ W; EPA, 18 Oct 1968. MZUSP 27331 (1, 104.4 mm SL): Tabatinga, rio Solimões, Lago Caial, c. 4 ° 21 ’ S, 69 ° 52 ’ W; L. P. S. Portugal, 8 Oct 1982. Amazonas state, rio Uatumã basin: INPA 16409 (1, 184.1 mm SL): igarapé Água Branca, trib. rio Pitinga (trib. rio Uatumã), c. 0 ° 52 ’ S, 59 ° 27 ’ W; Eq. Ictiologia / INPA, 17 April 1983. INPA 16355 (2, 194.6 – 267,0 mm CP), Balbina, cachoeira do Boto; Rômulo & Walter, 29 Oct 1987. INPA 5510 (1, 236.4 mm SL): rio Uatumã, Água Branca; S. Amadio, May 1985. Amazonas state, rio Negro basin: INPA 15730 (1, 174.2 mm SL): rio Tarumã-Açú, igarapé Tarumãzinho, BR- 174, km 28, 2 ° 43 ’ 51 ’’ S, 60 ° 4 ’ 88 ’’ W; A. Kirovsky, 10 Sept 1993. MZUSP 6191 (3, 192.7 – 208.9 mm SL): igarapé Jaraqui, trib. rio Negro, above Manaus; EPA, 22 – 24 April 1967. MZUSP 56810 (1, 148.4 mm SL): Anavilhanas, Lago do Prato, Paraná, c. 2 ° 45 ’ S, 60 ° 48 ’ W; G. Borges, Sept 1981. MZUSP 57032 (1, 144.1 mm SL); MZUSP 57031 (13, 230.1 – 274.9 mm SL): Anavilhanas, rio Negro, c. 2 ° 51 ’ S, 60 ° 38 ’ W; G. Borges, May 1982. MZUSP 56783 (1, 228.3 mm SL): rio Negro, Anavilhanas (igapó), c. 2 ° 51 ’ S, 60 ° 38 ’ W; M. Goulding, May 1980. MZUSP 56786 (2, 144.6 – 161.7 mm SL): Anavilhanas, igapó, c. 2 ° 51 ’ S, 60 ° 38 ’ W; G. Borges, March 1981. INPA 16378 (2, 163.8 – 170.9 mm SL): rio Negro, Ponta da Piraíba; Eq. Ictiologia / INPA, 15 Aug 1985. Amazonas state, rio Madeira basin: MZUSP 58693 (3, 120.0 – 140.3 mm SL): rio Canumã, c. 4 ° 2 ’ S, 59 ° 6 ’ W; EPA, 28 – 29 Nov 1967. Amazonas state, rio Juruá basin: AMNH 12553 (1, 210.2 mm SL): mouth of rio Envira (trib. Rio Tarauacá), near Envira, 7 ° 30 ’ S, 70 ° 4 ’ W; B. A. Krukoff, 1935. Rondônia state, rio Madeira basin: MZUSP 14027 (2, 219.2 – 273.1 mm SL): Lago Cururu, rio Machado, c. 8 ° 4 ’ S, 62 ° 53 ’ W; M. Goulding, April 1978. INPA 16446 (3, 202.2 – 247.7 mm SL): rio Jamari, below Samuel dam (igapó), c. 8 ° 38 ’ S, 63 ° 31 ’ W; G. M. Santos, 27 March 1986. INPA 16441 (1, 289.1 mm SL); INPA 16445 (1, 234.0 mm SL): rio Jamari, above Samuel dam, c. 9 ° 30 ’ S, 63 ° 7 ’ W; G. M. Santos, Dec 1984. INPA 16360 (1, 149.3 mm SL): rio Jamari, c. 5 km above Samuel dam; G. M. Santos, 14 June 1988. INPA 16434 (1, 200.8 mm SL): rio Machado, c. 20 km below Ji-Paraná, c. 10 ° 46 ’ S, 61 ° 54 ’ W; G. M. Santos, 3 Jul 1984. Mato Grosso state, rio Madeira basin: MZUSP 60398 (1, 146.0 mm SL): Aripuanã, rio Aripuanã, ferry on road Colniza / Panelas, 9 ° 34 ’ 45 ’’ S, 59 ° 25 ’ 19 ’’ W; F. A. Machado et al., 22 – 23 July 1997. MZUSP 77403 (1, 128.8 mm SL): Panelas, rio Roosevelt, above falls, 9 ° 11 ’ S, 60 ° 44 ’ W; F. A. Machado et al., 17 – 18 July 1997. Roraima state, rio Branco basin: MZUSP 56779 (4, 204.7 – 227.9 mm SL); MZUSP 31447 (1, 234.1 mm SL): Caracaraí, rio Branco, cachoeira do Bem Querer, tributary 2 km above rapids, 1 ° 58 ’ N, 61 ° 0 ’ W; M. Goulding, Jan 1984. Pará state, rio Trombetas basin: MZUSP 5455 (1, 155.2 mm SL); MZUSP 58651 (1, 132.7 mm SL): Oriximiná, rio Trombetas, 1 ° 45 ’ S, 55 ° 52 ’ W; EPA, Dec 1967. MZUSP 56780 (1, 167.8 mm SL): rio Trombetas, Cuminá, c. 1 ° 29 ’ S, 56 ° 1 ’ W; M. Goulding, Oct – Nov 1983. INPA 5566 (1, 224.5 mm SL): rio Trombetas, Lago Abuí, 1 ° 16 ’ S, 56 ° 57 ’ W; E. Ferreira & J. Zuanon, 1 Sept 1990. Pará state, rio Amazonas basin: MPEG 14124 (1, 163.1 mm SL): Juruti, rio Amazonas, lago Jará, 2 ° 9 ’ 19 ’’ S, 56 ° 4 ’ 6 ’’ W; L. F. A. Montag, 1 Dec 2007. MPEG 13977 (2, 156.6 – 158.0 mm SL): Juruti, rio Juruti Grande, 2 ° 22 ’ 55,1 ’’ S, 56 ° 18 ’ 51.6 ’’ W; L. F. A. Montag, 27 Nov 2007. MZUSP 3574 (3, 127.5 – 141.9 mm SL): Santarém, rio Tapajós, c. 2 ° 25 ’ S, 54 ° 45 ’ W; A. Amaral Campos, Oct 1944. MNHN 1909 – 070 (1, 115.1 mm SL): Santarém, “ fleuve Amazone ”, c. 2 ° 25 ’ S, 54 ° 45 ’ W; C. Jobert, no date. MCZ uncat. (3, 255.0 – 295.0 mm SL): Pará, rio Tapajós [no precise locality]; N. Dexter & W. James, 26 Aug 1865. Acre state, rio Juruá basin: ZUEC 6800 (1, 223.0 mm SL): Cruzeiro do Sul, mouth of rio Moá, 7 ° 39 ’ 28 ’’ S, 72 ° 40 ’ 38 ’’ W; G. V. Andrade, 2 June 1982. Peru, Depto. Loreto, rio Amazonas basin: MZUSP 15267 (3, 51.1 – 67.0 mm SL): Santa Maria Hojeal, Río Amazonas; M. V. Correa, 26 May 1980. CAS 136581 (1, 82.3 mm SL); CAS 160517 (1, 88.7 mm SL); CAS 160753 (1, 118.2 mm SL); CAS 53426 (1, 92.0 mm SL); CAS 136580 (1, 89.2 mm SL); CAS 136579 (2, 127.3 – 127.6 mm SL): Cano del Shansho, near Pebas, c. 3 ° 21 ’ S, 71 ° 47 ’ W; W. G. Scherer, 1936 – 1937. CAS 117266 (1, 95.3 mm SL); CAS 160574 (1, 90.0 mm SL): Rio Ampyiacu, near Pebas, c. 3 ° 21 ’ S, 71 ° 47 ’ W; W. G. Scherer, 1937 – 1940. MNRJ 4008 (1, 100.2 mm SL): near Pevas (= Pebas, c. 3 ° 21 ’ S, 71 ° 47 ’ W); W. G. Scherer, 1941. USNM 175983 (2, 141.7 – 156.3 mm SL): Tuye Caño, Río Ampyiacu drainage; W. G. Scherer, 17 Aug 1935. NRM 23399 (1, 107.3 mm SL): just below Esperanza village, mouth area of small quebrada tributary to Río Yaguasyacu, Río Ambyacu drainage, c. 3 ° 21 ’ S, 71 ° 47 ’ W; S. O. Kullander et al., 12 Aug 1981. USNM 280505 (1, 110.8 mm SL): caño entering Río Manite 10 km upriver its mouth on Río Amazonas, 3 ° 32 ’ S, 72 ° 40 ’ W; R. P. Vari et al., 21 Aug 1986. USNM 280051 (1, 126.2 mm SL): Río Nanay at Nanay beach, 3 ° 50 ’ S, 73 ° 11 ’ W; R. P. Vari et al., 17 Aug 1986. MUSM 60 (2, 116.6 – 129.3 mm SL): Iquitos, Río Nanay, Porto Almondras; H. Ortega & A. Urteaga, 6 Jul 1984. NRM 23668 (1, 113.8 mm SL): Quebrada Corrientillo, 20 km from Iquitos on road to Puerto Almenda, c. 3 ° 50 ’ S, 73 ° 13 ’ W; S. O Kullander et al., 18 Aug 1983. MUSM 1978 (6, 112.3 – 117.7 mm SL): Iquitos, Quebrada Corrientillo, at Corrientillo, road Iquitos to Río Nanay, 3 ° 49 ’ 57 ’’ S, 73 ° 21 ’ 43 ’’ W; R. P. Vari et al., 19 Aug 1986. INHS uncat. (1, 102.8 mm SL); INHS 44000 (4, 93.5 – 119.2 mm SL): Río Nanay, Pampa Chica, 3 ° 45 ’ 1 ’’ S, 73 ° 17 ’ 0 ’’ W; M. H. Sabaj et al., 22 – 27 Jul 1997. ANSP 136819 (3, 97.5 – 116.4 mm SL): Iquitos, Río Nanay just above Cocha Moronona (c. 9 miles above mouth at Río Amazonas); M. Holm, 17 Oct 1955. ANSP 178408 (1, 128.2 mm SL): Río Amazonas, sandy beach upstream mouth of Río Yanuyacu, 4 ° 14 ’ 1 ’’ S, 73 ° 19 ’ 14 ’’ W; M. H. Sabaj et al., 11 Aug 2001. NRM 23397 (4, 104.3 – 107.0 mm SL): Varillal pool, Río Itaya drainage, c. 4 ° 13 ’ S, 73 ° 29 ’ W; S. O. Kullander et al., 4 Jul 1986. NRM 23673 (1, 99.3 mm SL): Quebrada Pintoyacu, where crossed by carretera Iquitos-Nauta; S. O. Kullander et al., 3 July 1986. NRM 23398 (2, 95.6 – 105.1 mm SL): pools near Quebrada Tocón Grande at km 33 on carretera Iquitos-Nauta, Río Itaya drainage, c .. 4 ° 2 ’ S, 73 ° 26 ’ W; S. O. Kullander et al., 3 July 1986. MUSM 34 (1, 109.4 mm SL): Hucuyhuasi Shante, Iquitos; F. Benitez, 19 May 1975. MUSM 12744 (5, 109.2 – 111.4 mm SL): Yanayacu, Base 3, Pluripetrol, Río Shiriyacu, 4 ° 51 ’ S, 74 ° 56 ’ W; H. Ortega, 10 Oct 1997. MUSM 18388 (2, 226.4 – 226.9 mm SL): Río Paztaza, near mouth of Quebrada Asnagra, 3 ° 17 ’ 23 ’’ S, 76 ° 23 ’ 3 ’’ W; H. Ortega et al., 5 Aug 1999. NRM 23395 (1, 114.7 mm SL): Base Tacsha, left bank sand playa of Río Samíria, c. 4 ° 41 ’ S, 74 ° 20 ’ W; S. O. Kullander et al., 8 Aug 1986. CAS 69078 (ex IU 16021) (2, 143.8 – 146.9 mm SL): Yarinococha (= Llarinococha), shallow, clear cutoff lake connected to Rio Pacaya by a long, narrow channel, c. 5 ° 9 ’ S, 74 ° 11 ’ W; W. R. Allen, 1 Sept 1920. MUSM 18268 (1, 122.0 mm SL): Cocha San Pablo, Tipishca, Comunidade Sucre; H. Ortega et al., 15 April 2001. MUSM 10892 (1, 86.8 mm SL): Río Chambira, Comunidad San Juan, c. 4 ° 36 ’ S, 74 ° 52 ’ W; E. Castro, 14 March 1997. MUSM 2660 (1, 117.3 mm SL): Iquitos, Tipishca del Río Samíria, c. 4 ° 52 ’ S, 74 ° 22 ’ W; C. Riofrio et al., 23 Nov 1989. USNM 167787 (2, 145.0 – 159.2 mm SL): mouth of Río Pacaya at Bretaña, c. 5 ° 13 ’ S, 74 ° 18 ’ W; W. R. Allen, Jul 1920. USNM 167789 (2, 125.9 – 151.2 mm SL): Río Ucayali, Orellana, 6 ° 55 ’ S, 65 ° 9 ’ W; W. R. Allen, Aug 1920. Depto. Ucayali: MZUSP 26644 (2, 128.1 – 138.2 mm SL): Ucayali, Pucallpa, Ivita (stocked from specimens collected at Iquitos); H. Ortega, 9 Sept 1976. MUSM 22 (3, 56.6 – 88.5 mm SL): same locality data as previous; H. Ortega, 28 Sept 1972. MUSM 3572 (1, 125.7 mm SL): Pucallpa, IVITA, Río Neshuya, Campo Verde; C. Rojas & G. Contreras, 24 Dec 1981. MUSM 53 (1, 159.3 mm SL): Pucallpa, Patria Nueva, Río Calleria near mouth of Río Ucayali, 8 ° 31 ’ S, 74 ° 35 ’ W; H. Ortega, 4 Oct 1984. AMNH 43314 (2, 61.1 – 62.8 mm SL); MUSM 2377 (2, 59.7 – 68.3 mm SL): Río Ucayali, Masisea, 8 ° 35 ’ S, 74 ° 20 ’ W; H. Ortega, 20 – 23 Nov 1973. Depto. Pasco: USNM 167788 (1, 163.4 mm SL): Río Pichis (trib. Río Pachítea), Puerto Bermudez, c. 10 ° 18 ’ S, 74 ° 56 ’ W; W. R. Allen, Jul 1920. USNM 167790 (2, 138.5 – 187.2 mm SL): Río Pachitea; W. R. Allen, Jul 1920. Depto. Madre de Dios: USNM 263993 (1, 269.9 mm SL): stream trib. Río Madre de Dios, 10 km below mouth of Río Tambopata, c. 12 ° 30 ’ S, 69 ° 10 ’ W; R. P. Vari et al., 25 Aug 1983. MUSM 9654 (1, 184.8 mm SL): Tambopata, Lago Valencia; C. Cañas, 19 June 1996. Peru, no precise locality: ANSP 21972 (3, 65.5 – 74.4 mm SL): “ Peruvian Amazon ”; J. Orton, no date. Ecuador: MEPN uncat. (1, 211.0 mm SL): Pastaza, Río Lliquino, 2 km SE Villano, 01 ° 24 ' S 77 ° 40 ' W; R. Barriga & J. Llaney, 2 Sept 1997. MCZ 49158 (1, 242.7 mm SL): Napo, Río Napo at Coca, 0 ° 28 ’ S, 76 ° 56 ’ W; T. R. Roberts, 28 Nov 1971. FMNH 102253 (1, 243.0 mm SL): Napo, Río Lagartococha, 1 km upstream its mouth at Río Aguarico, 0 ° 38 ’ S, 75 ° 15 ’ W; D. J. Stewart et al., 1 Nov 1983. FMNH 102252 (1, 148.5 mm SL): Napo, Río Tiputini, near mouth at Río Napo, 0 ° 49 ’ S, 75 ° 31 ’ W; D. J. Stewart et al., 30 Oct 1981. FMNH 103394 (1, 293.0 mm SL): Laguna Grande de Cuyabeno and Quebrada de Hormigas, Río Aguarico drainage, 0 ° 0 ’ 30 ’’ S, 76 ° 11 ’ 30 ’’ W; D. J. Stewart et al., 28 Sept 1983. MEPN 10926 (1, 244.0 mm SL): Orellana, tributary of Río Tiputini, Pozo Zine; no collector / date specified. MEPN 11122 (1, 228.0 mm SL): Sucumbios, Río Napo at Garzacocha, 0 ° 30 ’ S, 76 ° 22 ’ W; no collector / date specified.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FFAFF5B4EA4F900FF22FAB2.taxon	description	Brycon sp. (B. falcatus group): Toledo-Piza, 2002: 116 – 117 (drawing by A. R. Wallace; “ Rio Uaupés ”; common name). Brycon carpophagus (not Valenciennes): Mérona et al., 2001: 387, 391 (Rio Tocantins, Tucuruí; diet before and after river impoundment).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FFAFF5B4EA4F900FF22FAB2.taxon	diagnosis	Diagnosis. Brycon falcatus can be distinguished from all remaining cis-andean Brycon species, with the exception of B. gouldingi, by typically possessing a V- or crescent-shaped dark blotch on caudal peduncle and caudal fin. Brycon falcatus can be distinguished from B. gouldingi by possessing possessing dark, longitudinal stripes formed by pigmentation concentrated on the mid-distal portion of scales (vs. dark, wavy longitudinal stripes formed by dark pigment concentrated on upper and lower scale margins), paired fins typically clear (vs. paired fins black), and lower lateral line scale counts (47 – 69, modally 57, vs. 66 – 82, modally 74). Brycon falcatus is morphologically more similar to B. melanopterus, and an unequivocal diagnosis between both species based only on external morphological characters is not possible due to the great polymorphism in color pattern observed in the species. However, typically, Brycon falcatus does not present an oblique dark solid stripe, presenting instead a V or crescent-shaped blotch on caudal fin (vs. black pigmentation restricted to the upper caudal-fin lobe in B. melanopterus). In addition, Brycon falcatus is typically a higher bodied fish, with higher vertebrae counts. See the item “ Comparisons ” under Brycon melanopterus, and the item “ Variation ”, below, for thorough account on the subject.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FFAFF5B4EA4F900FF22FAB2.taxon	description	Description. Morphometric data are presented in Tables 21 – 22; some meristic data (lateral line counts and scales between dorsal-fin origin and lateral line) in Tables 23 – 24. Middle- to large-sized species, largest examined specimen 434.7 mm SL. Body moderately slender to 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 moderately 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 to pronouncedly 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, snout obtuse, 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. Five (2), 6 (5), 7 (29), 8 (65), 9 (118), 10 (99), 11 (53), 12 (8), or 13 (1) relatively small tricuspidate teeth in outer series. Three (30), 4 (229), 5 (107), 6 (6), or 7 (1) tri- to pentacuspidate teeth in second, inner premaxillary row, plus 2 (2), 3 (150), 4 (218), or 5 (7) 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, both pentacuspidate. Maxillary margins approximately parallel, straight in profile. Fifteen to 28 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 7 (7), 8 (34), 9 (72), 10 (61), 11 (26), 12 (9), 13 (3), 14 (1), or 15 (1) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, pentacuspidate, 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 13 – 21 small, aciculated, unicuspidate teeth, originating on lingual crest of dentary replacement trench at the level of fourth to seventh main series dentary teeth. Symphyseal teeth may be lacking in larger (> 350 mm SL) specimens. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-seven (1), 48 (1), 49 (1), 50 (3), 51 (11), 52 (25), 53 (57), 54 (59), 55 (77), 56 (90), 57 (104), 58 (93), 59 (57), 60 (49), 61 (27), 62 (14), 63 (8), 64 (16), 67 (1), or 69 (1) 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 (> 270 mm SL) specimens. Horizontal scale rows between dorsal-fin origin and lateral line eight (1), 9 (21), 10 (115), 11 (122), 12 (103), 13 (29), or 14 (3). Horizontal scale rows between lateral line and pelvic-fin 4 (21), 5 (156), 6 (169), 7 (27), or 8 (1). Circumpeduncular scales 15 (1), 16 (27), 17 (75), 18 (125), 19 (67), 20 (51), 21 (32), or 22 (6). Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead, or about at middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 11 th (2), 12 th (1), or 13 th (1) vertebra. Anal-fin rays iii (not including first, small unbranched ray only visible in cs specimens), 18 (1), 19 (4), 20 (11), 21 (41), 22 (90), 23 (93), 24 (82), 25 (45), or 26 (6). First anal-fin pterygiophore inserting behind haemal spine of 22 th (4), or 23 th (1) vertebrae. Anal-fin rays decreasing only slightly in size towards anal-fin end. Anal fin displaying several (c. 5 – 12 per fin-ray main branch) small hooks on posterior main branch of branched rays 2 – 11, associated with dense, gelatinous tissue in 5 examined specimens (MZUSP 3845, 2, 196.2 – 198.9 mm SL, MZUSP 16448, 1, 196.5 mm SL mm SL, MZUSP 91493, 1, 226.0 mm SL, MZUSP 94990, 1, 230.0 mm SL). A single hook per ray segment. Sheath of scales covering basis of anal-fin rays composed of four scale rows, lower scale row formed by 21 – 25 rectangular scales. Pectoral-fin rays 11 (3), 12 (76), 13 (229), or 14 (58). Pelvic-fin rays typically i, 7 (364), rarely i, 6 (4), or i, 8 (4). Main caudal-fin rays 10 / 9. Caudal fin forked, lobes slightly emarginated. 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. Small distal projection in the area of the laterosensory tube present in specimens with relatively intact caudal fin. Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First branchial arch with 12 (14), 13 (24), 14 (40), 15 (32), 16 (22), or 17 (2) lower, 1 at angle, and 11 (1), 12 (15), 13 (38), 14 (51), 15 (20), 16 (6), or 17 (2) upper gill rakers. Vertebrae 39 (1), 40 (1), 41 (7), 42 (5), 43 (2), or 44 (1). Supraneurals 8 (2), or 9 (2). Coloration in alcohol. Top of head, snout, supraorbital, and sixth infraorbital light-grey to light-brown. Dorsal portion of body light-grey to dark-brown. Second, third, fourth, and fifth infraorbitals, opercle and cleithrum silvery in specimens retaining 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 silvery in specimens retaining guanine, light brown in specimens that lost this pigment due to a long storage in formalin. Humeral blotch present, slightly to moderately conspicuous, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of second, extending longitudinally to posterior margin of fourth to fifth lateral line scales, and vertically one and half scales high. Scales on lateral portions of body with dark pigmentation concentrated on their central portion, forming dark, straight longitudinal stripes, more conspicuous dorsally. Dark, solid, oblique dark stripe extending along anal-fin scales sheath and anal-fin basis in almost all specimens (see item “ Variation ”, below). Caudal peduncle blotch wide, extending across upper and lower caudalfin rays typically as a V- or crescent-shaped dark blotch, though with a great degree of variation in the development of the pigmentantion in the lower caudal-fin lobe (see item “ Variation ”, below). Dorsal, anal, pectoral, and pelvicfin rays with a variable amount of interradial dark pigmentantion but typically clear. Adipose-fin light- to darkgrey, with scattered dark spots in large specimens from the upper rio Tapajós basin. Color in life. Description based on pictures of several specimens collected at the lower rio Japurá (Lago Amanã), upper rio Madeira, upper rio Tapajós, upper rio Xingu (Fig. 90), and rio Tocantins in Amazon basin, Río Inirida, Orinoco basin, and Kuribrong River, Essequibo basin, plus published pictures of living specimens (Géry, 1977: 321, upper and lower; Planquette et al., 1996: 227; Ferreira et al., 2007: 125). Top of head and snout light- to dark-grey. Dorsum plumbeous. Lateral surfaces of head and body silvery. Humeral blotch relatively inconspicuous. Dark stripe extending along anal-fin basis and dark, V-shaped caudal peduncle / caudal-fin blotch very conspicuous. Distal border of caudal fin yellowish to reddish. Adipose fin yellowish grey. Remaining fins translucent, dorsal fin with some yellowish pigmentation. Variation. Brycon falcatus is one of the most widespread cis-andean Brycon species, and perharps not unexpextedly, a great degree of variation in body shape and color pattern is observed across its range. Specific features that show a great degree of variation are body height, caudal-fin pigmentation, scale counts, maximum body size, and, to a lesser extent than the previous features, anal-fin pigmentation. Each of these features is treated sequentially below. Body height varies considerably in specimens of Brycon falcatus, to an extent unequalled among cis-andean Brycon species. Specimens range from being relatively elongated (Fig. 89) to high-bodied (Figs. 86, 88, 92). However, there is no consistent geographical variation in this character, and most river systems present specimens ranging from both extremes in body shape. Probably, most of the observed variation in this trait is simply a result of phenotypic plasticity. Caudal-fin pigmentation is the most significant trait to vary among Brycon falcatus populations, and, contrary to body height, exhibits a clear geographical pattern. Brycon falcatus populations from the Guianas, rio Branco, rio Capim, rio Tocantins, and rio Xingu basins possess the characteristic V- or crescent-shaped caudal-fin blotch, with the extent of dark pigmentation approximately symmetrical in both lobes (Figs. 86 – 87). However, moving westwards into the Brazilian and Guyanese shields, dark pigmentantion on the lower caudal-fin lobe tend to decrease clinally in intensity. Specimens from the rio Tapajós and rio Madeira basins range from possessing an almost symmetrical dark pigmentation in both fin lobes (Figs. 88, 92) to a considerably less pigmented lower caudal-fin lobe. Specimens from the middle and lower rio Negro and middle and middle rio Orinoco basin possess a clearly asymmetrically pigmented caudal fin, with the dark pigmentation much more intense and extensive in the upper, than in the lower caudal-fin lobe (Figs. 93 – 95). Moving westward into western Amazon, the apparently scattered populations of Brycon falcatus occurring in that vast area possess an even more asymmetrically pigmented caudal-fin, with the lower lobe almost (Fig. 96) or completely (Fig. 97) unpigmented, the latter condition found in specimens from Iquitos area, the most westward known locality for the species. Oddly, specimens from the upper rio Negro (above São Gabriel rapids) and upper rio Orinoco (above its junction with the Río Guaviare) basins present a similar color pattern to the specimens from Iquitos area, presenting no dark pigmentation at all at the lower caudal-fin lobe and the upper caudal-fin lobe pigmentation almost continuous with the anal-fin dark stripe (Fig. 98). Specimens from the Iquitos area and upper rio Negro and upper Río Orinoco basins are thus very similar in color pattern to B. melanopterus, though B. falcatus is typically a higher-bodied species, with lower vertebral counts (see item “ Comparisons ” of B. melanopterus). See item “ Putative examples of mimicry involving Brycon ”, below, for a possible explanation for this similarity. Scale counts vary considerably among distinct populations of Brycon falcatus, with specimens from the Guianas clearly presenting lower counts when compared with specimens from the Orinoco and Amazon basins (see Tables 23 and 24). The lower scale counts presented by Guyanese specimens when compared with specimens from rio Capim and rio Tocantins basins were the main reason supporting the recognition of Brycon brevicauda as a valid species (e. g., Géry, 1964: 450; Howes, 1982: 14). However, the examination of large series of specimens undertook during the present study demonstrated a continuous variation in scale counts among the several populations of Brycon falcatus, thus failing to confirm the presumed distinctness of B. brevicauda based on this character (Tables 23 and 24). Another trait which shows a clear geographical component in Brycon falcatus is the maximum length reached by different populations. Typically, Brycon falcatus is a middle-sized species, generally does not exceeding 300 mm SL (e. g., the maximum size of 350 mm TL reported by Rubio et al., 2012, based on the examination of 279 specimens collected at the rio Guaporé). However, extensive examination of both preserved and unpreserved specimens of Brycon falcatus from the rio Teles Pires and rio Juruena in the upper rio Tapajós basin, and from the upper rio Xingu basin showed that individuals belonging to these populations often grow considerably above this upper limit. Maximum recorded sizes from specimens from these areas are, respectively, 435.0 and 401.0 mm SL, and specimens ranging between 300 – 400 mm SL are common. To complicate further the issue, in the upper rio Tapajós area apparently both morphotypes (“ large-sized ” and “ middle-sized ”) co-exist (J. M. Mendes and H. F. Mendes, pers. comm.). The “ large-sized ” population from the upper rio Xingu is slightly different from the one from the upper rio Tapajós in being on average smaller-sized and by not possessing the dotted adipose fin present in the latter. 58 59 60 61 62 63 64 65 66 67 68 69 Guyana and Suriname Orinoco 4 1 1 1 Negro 6 2 1 Branco 4 2 Tocantins and Capim 11 12 11 8 2 2 5 Xingu 7 4 2 1 Tapajós 16 10 10 6 2 6 2 Madeira 2 2 1 3 1 Western Amazon 1 1 1 1 8 9 10 11 12 13 14 Guyana and Suriname 1 22 33 Orinoco 10 7 1 6 Negro 14 24 1 Branco 12 18 Tocantins and Capim 12 44 4 Xingu 8 11 21 8 Tapajós 2 28 23 25 7 Madeira 8 16 11 Western Amazon 4 11 1 2 Brycon falcatus typically possess a conspicuous dark stripe along the basis of anal-fin, the intensity of which, however, varies considerably among populations and even within specimens collected in the same general area. Some specimens from the upper portion of the rio Araguaia in central Brazil (MZUSP 18628, MZUSP 62490, MZUSP 48113, MZUSP 52391) entirely lack the dark stripe. At the other extreme are the specimens from the upper Río Orinoco and upper rio Negro, which possess a very well-developed dark stripe that extends from the origin of the pelvic fin to the caudal peduncle, much as the stripe found in Brycon melanopterus (see above). It is interesting to notice that specimens collected in the rio Arinos at the upper rio Tapajós basin (e. g., MZUSP 56785, MZUSP 60423, MZUSP 61056) possess a well-developed dark stripe, while specimens collected in the same area but kept for some time in fish ponds (MZUSP 61132, MZUSP 61133) lack entirely the dark stripe. This suggests that the expression and the intensity of the dark stripe might vary as a response of some physiological factor. Finally, a considerable molecular divergence among samples the species from the Río Orinoco in Venezuela, rio Negro in Brazil and upper rio Xingu was detected by Abe et al. (2014). Abe et al. (2014: 12) considered that populations of Brycon falcatus from the Río Orinoco and Río Negro basins were separated by the development of the Vaupes arch. However, Brycon falcatus possess a continuous distribution across the upper rio Negro and upper Río Orinoco afforded by the Río Casiquiare, from where the species is known (see Fig. 99), and consequently the genetic divergence between these samples cannot be ascribed to a supposed vicariant event that in fact did not isolated these populations. A much larger sampling is necessary to understand phylogeographical patterns and the possible existence of cryptic species within the populations herein assigned to Brycon falcatus. Until such work is done, it is deemed more parsimonious to consider all these populations as belonging to a single species. Common names. Brazil: “ matrinchã ”; “ piabanha ” (Santos et al., 1984: 41); “ voadeira ”, “ avoadeira ”, “ matrinchã-pequena ”; “ jatuarana-miri ” (Wallace, in Toledo-Piza, 2002: 116); “ miõ wi ” (Tukano language, upper rio Negro basin; Lima et al., 2005: 164); “ wenawe ” (Tuyuka language, upper rio Negro basin; Lima et al., 2005: 164); French Guiana: “ moloko blanc ”, “ maloko ”, “ mbooko ”, “ molokoimo ” (Planquette et al., 1996); Suriname: “ moroko ” (Richter & Nijssen, 1980: 123; Mol et al., 2007: 115); Guyana: “ kurumi ” (Müller & Troschel, 1845: 29); Venezuela: “ palambra ”, “ bócon ” (Taphorn, 1992: 128).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FFAFF5B4EA4F900FF22FAB2.taxon	distribution	Distribution. Widespread in rivers draining the Brazilian and Guyana shields in the Amazon, Orinoco, and guyanese river systems, and also on scattered localities in western Amazon (Fig. 99). Brycon falcatus is recorded only from clear or dark water rivers and never occur at white / murky-water rivers. Records from the rio Madeira basin are all from shield-draining, clear- to black-water tributaries such as rio Guaporé / Itenez, rio Jamari, rio Aripuanã, and rio Roosevelt. Likewise, records in the western Amazon came from black-water systems as the Lago Amanã near the mouth of rio Japurá in Brazil or the Río Nanay near Iquitos, Peru. From guyanese river systems, we have examined material from the Essequibo, Corantijn, and Maroni / Marowijne Rivers in Guyana, Suriname and French Guiana, but the species is also reported from the Suriname River (e. g., Richter & Nijssen, 1980; Mol et al., 2007; Mol et al., 2012), Coppename River (Mol et al., 2006; Mol et al., 2012), Nickerie River (Mol et al., 2012) and Saramacca River (Mol et al., 2012) in Suriname, and Mana River (Planquette et al., 1996: 226 – 227) in French Guiana. The present westernmost record for Brycon falcatus is Iquitos and records from more westward localities (e. g., Ecuador; Gilbert & Roberts, 1971; Saul, 1975) need to be re-examined. Recorded as a vagrant species in a river from southern Trinidad (Phillip et al., 2013: 16; see item “ Biogeography ”, below). Ecological notes. As noticed above, Brycon falcatus inhabits exclusively clear- or black-water rivers. Like its congeners, Brycon falcatus is a highly mobile fish and utilizes a great variety of habitats, ranging from river channels, flooded forest, floodplain lakes, to small tributaries. The species undertake massive spawning upstream migrations in the rio Culuene, upper rio Xingu basin, during September / November (pers. obs.), but spawning migrations in the rio Araguaia basin appear to be much more discrete and local (J. B. Nunes, pers. comm.). Specimens associated in small groups of up to 3 individuals were observed while snorkelling in clear-water tributaries of the upper rio Arinos, rio Tapajós basin, Mato Grosso during the late wet season (pers. obs.). Lowe Mc-Connel (1964: 118, 120) noticed downstream migration of the species in the Ireng River (upper Rio Branco basin, Guyana) during the early dry season, and specimens with ripening gonads in the Rupununi River in April (early wet season). At the rio Guaporé (Mato Grosso, Brazil), mature females were collected at the beginning of the rainy season, between October and December (Rubio et al., 2012). Scale rings indicated a maximum age of five years in specimens from the rio Guaporé (Rubio et al., 2012). A female with 30 cm SL from the rio Culuene collected early October possessed mature gonads (pers. obs.). Migratory schools in the rio Culuene were observed being attacked by Hydrolycus armatus specimens, and some dying Brycon falcatus were found with deep parallel cuts which very likely were inflicted by this large predatory characin (pers. obs.). An osprey, Pandion haliaetus, was observed with a Brycon falcatus specimen under its talons at the Kuribrong River, Guyana (pers. obs.). Detailed studies on the diet of Brycon falcatus are Borges (1986), who studied specimens from the middle and lower rio Negro basin in Brazil, and Albrecht et al. (2009), studying specimens from the upper rio Tocantins basin in central Brazil. Borges (1986) found mainly vegetal itens (fruits and seeds) in stomach contents of specimens collected during the rainy season, while during the dry season, arthropods were the main dietary item. Leaves, flowers, and fishes were moderately consumed in both seasons. Fruits commonly found in stomach contents were Ocotea sp. (Lauraceae), Tococa sp. (Melastomataceae), and Macrolobium acacifolium (Fabaceae). Albrecht et al. (2009) also found seeds and fruits to be the most important dietary item for the species, followed by ants, coleopterans, fish and terrestrial vertebrates. Correa & Winemiller (2014) reported that the species switched from fruits and seeds to terrestrial invertebrates as its main dietary item from rainy season to dry season in a site at the lower Río Caquetá basin (= Rio Japurá in Brazil) in Colombia, although later than the syntopic B. melanopterus. Matos et al. (2016) recorded invertebrates (mostly insects and Decapoda), fishes, flowers, seeds and fruits as the main itens ingested by Brycon falcatus at the rio Teles Pires basin, and that specimens living in river stretches where an artificial food supply (soybeans and corn) was provided presented a better condition (i. e., more abdominal fat) than specimens living in river stretches where no artificial food supply was provided.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FFAFF5B4EA4F900FF22FAB2.taxon	discussion	Remarks. Jardine, in Jardine & Schomburgk (1841: 212 – 213) described Chalceus labrosus from the “ river Paduiri ” (= Rio Padauari), a locality which lies in the middle rio Negro basin, Amazonas, Brazil, an area visited by Schomburgk in March 1839 (Schomburgk, 1840). No type material is known for this nominal species. Though poor, the description and plate of Chalceus labrosus clearly correspond to the same species latter described by Müller & Troschel as Brycon falcatus. Jardine, in Jardine & Schomburgk (1841: 212 – 213) mentions the three series of teeth in the premaxillary (“ three rows on the roof of the mouth ”), the dark stripe at the basis of anal fin (“ a broad black mark running from the centre of its anterior edge for one-third of its length ”), and the crescent-shaped blotch at the caudal fin (“ a dark bluish black bar running across, parallel to the fork ”). The name Chalceus labrosus was never used as a valid species after its original description, while Brycon falcatus has been continuously used in the ichthyological literature during the last 160 years, by well more than 25 authors (see synonymic list, above). We consider thus that Chalceus labrosus fulfills the necessary requisites to be considered as a nomen oblitum (Article 23.9.1 of the International Code of Zoological Nomenclature, 1999), and, as such, as not having precedence over Brycon falcatus, which is herein considered a nomen protectum. Müller & Troschel (1844: 90) briefly described Brycon falcatus, based on an unspecified number of syntypes collected at Guiana and Suriname by Schomburgk and Dieppering. A little later, the same authors (Müller & Troschel, 1845: 29, pl. 6, fig. 1) described the species in detail and provided a good illustration of it. Though the type specimens were not examined in the present study, the good description and illustration by Müller & Troschel (1845) leaves no doubt as to the identity of the species. Günther (1864: 335) identified three specimens from BMNH collected by Schomburgk as belonging to the type series, which probably prompted Eschmeyer (1998: 58) in considering this lot (BMNH 1969.12.12: 1 - 3) as being part of the syntypical series of B. falcatus. However, as remarked earlier by Howes (1982: 26), only the specimens deposited at the ZMB were cited in the original description (Müller & Troschel, 1845: 29) and as such are the only specimens that should be considered as having typical status (Lima, 2003: 176). In the same papers where Brycon falcatus was described, Müller & Troschel (1844: 91; 1845: 29 – 30, pl. 6, fig. 2) described Brycon schomburgkii from Guyana. Müller & Troschel (1845) diagnosed the species from Brycon falcatus only by the possession of a longer inner row of dentary teeth, almost reaching the inner symphyseal teeth. However, there is a reasonable degree of intraspecific variation in the extension of the second dentary teeth row in Brycon species, and the variation described by Müller & Troschel (1845) fits within this variation. Eigenmann (1912: 372), who examined the type specimens of both Brycon falcatus and B. schomburgkii, remarked that “ the later is undoubtedly the young of the former ”, thus considering B. schomburgkii a synonym of B. falcatus, an opinion with which we concur. Valenciennes (in Cuvier & Valenciennes, 1850: 246 – 248), when describing Chalceus hilarii, mentioned that part of the syntypical series was collected “ dans l’Amazone ” by Castelnau. Castelnau (1855: 68) noticed that these specimens were actually collected at Salinas, located at the rio Araguaia basin in Goiás state, Brazil (Papavero, 1971: 152). Bertin (1948: 14 – 15) listed three syntypes for Chalceus hilarii, two of which (MNHN 9893, MNHN 9894) were collected by Castelnau at the “ Fl [euve]. Amazone ”. Géry & Mahnert (1992: 815) selected the specimen MNHN A. 8616 as the lectotype of Brycon hilarii (see item “ Remarks ” of B. hilarii). Consequently, the specimens MNHN 9893 and MNHN 9894 became paralectotypes of the species. Géry & Mahnert (1992) mentioned that these specimens were collected at Salinas, but incorrectly surmised that this locality was located at the rio Jequitinhonha (a river basin not visited by Castelnau). Due to the poor state of preservation of the paralectotypes, Géry & Mahnert (1992) were unsure whether they were in fact conspecific with the lectotype or not. The examination of one of these paralectotypes (MNHN A. 9894) demonstrated, as should be expected by its provenance, that it refers not to Brycon hilarii, but, actually, to B. falcatus. Günther (1864: 335) described Brycon brevicauda based on three syntypes, BMNH 1864.4.20: 13, from “ Rio Jocintins ” (presumably a mispelling of rio Tocantins), and BMNH 1849.4.8: 42; 49, from rio Capim, Brazil. Géry (1964: 450) provided an identification key for what he supposed to be a group of species similar to Brycon falcatus, identifiable by possessing a V-shaped caudal-fin blotch. In that key, Brycon brevicauda was diagnosed from B. falcatus by possessing higher lateral-line and vertical scale counts, lower anal-fin counts, a smaller body depth, by lacking the dark stripe at the anal-fin basis and, possibly, by lacking the inner dentary symphyseal teeth. Howes (1982: 14) also considered Brycon brevicauda as being a valid species, distinct from Brycon falcatus in scale counts, body depth, premaxillary teeth counts, and dentary teeth shape. The examination of large samples from the Guianas, Amazon and Orinoco basins undertook during the present study showed that all these characters are either not valid (i. e., premaxilary teeth counts, lack of symphyseal teeth, dentary teeth shape), or vary considerably within populations (i. e., body depth, anal-fin basis pigmentation), or else present a continuous variation among distinct populations (i. e., scale counts) (see item “ Variation ”, above). We consider thus Brycon brevicauda as a synonym of B. falcatus. Steindachner (1882 a: 176) briefly described Brycon stuebelii (originally spelled as stübelii) having as its type locality the Amazon basin (“ Amazonestrome ”). Soon thereafter Steindachner (1882 b: 13 – 14) redescribed the species in detail, and specified its type-locality as being “ Rio Amazonas (Iquitos) ”. Steindachner (1882 b: 13) compared Brycon stuebelii with a single congener, B. melanopterus, from which he diagnoses it by possessing a shorter dark stripe at the anal-fin basis and a supposedly larger second infraorbital bone. Géry (1964: 450) considered Brycon stuebelii as provisionally valid and considered it as belonging to a group of species similar to B. falcatus, being supposedly distinct from the latter by possessing higher scales counts. Howes (1982: 45), however, considered Brycon stuebelli as a possible synonym of B. falcatus. The holotype of Brycon stuebelii was not examined during the course of the present study, but Steindachner’s description and illustration show a typical Brycon falcatus specimen, with a well-defined crescent-shaped blotch on the caudal-fin, a color pattern no longer displayed by the now faded holotype (Zarske, 2003: 16, fig. 7). Curiously, all subsequent specimens of Brycon falcatus collected in the western portions of the Amazon basin, including specimens collected at Iquitos, does not possess a V- or crescent-shaped blotch on caudal fin, but instead, the dark pigmentation is considerably more developed or even entirely confined to the upper caudal-fin lobe (see item “ Variation ”, above).	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FFAFF5B4EA4F900FF22FAB2.taxon	description	Géry & Mahnert (1992: 800 – 802) identified as a possible new species specimens of Brycon belonging to the INPA collection from the rio Aripuanã and Lago Amanã. These specimens were considered by Géry & Mahnert (1992) to be distinct from Brycon brevicauda, and similar to B. bicolor, by possessing the dark pigmentation confined to the upper caudal-fin lobe. We have examined one of the lots studied by Géry & Mahnert (1992) (INPA 16412, from the rio Aripuanã) and specimens from the Lago Amanã and nearby areas (MZUSP 99213, MCP 29776, MCP 29774, MCP 29771, MCP 29775). As noticed under the item “ Variation ”, above, the intensity of pigmentation in the lower caudal-fin lobe vary in intensity across the populations of Brycon falcatus, with a distinct east-west clinal tendence of decrease of intensity of pigmentation in the lower caudal-fin lobe. The population of Brycon falcatus from the rio Madeira basin is variable in this respect, and specimens ranging from a lower caudalfin lobe much less pigmented than the upper caudal-fin lobe to specimens presenting a almost symmetric, V-shaped caudal-fin blotches are present in the area, while specimens from the lago Amanã area typically possess a lower caudal-fin lobe much less pigmented than the upper caudal-fin lobe, a condition which is in fact a rule for B. falcatus populations from the western Amazon (see item “ Variation ”, above). In sum, the purported specific distinctness of the populations from the rio Aripuanã and Lago Amanã hypothesized by Géry & Mahnert (1992) constitute actually only in the extreme of a continuous variation of the lower-caudal fin pigmentation within Brycon falcatus populations.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
C033D7104FFAFF5B4EA4F900FF22FAB2.taxon	materials_examined	Material examined. Type material. MNHN A. 9894 (1, 169.5 mm SL): " L'Amazone " (= Brazil, Goiás, Salinas, c. 13 ° 40 ’ S, 50 ° 14 ’ W, rio Araguaia basin); F. de Castelnau, 1844. Paralectotype of Chalceus hilarii Valenciennes (designated by Géry & Mahnert, 1992: 815). BMNH 1849.4.8: 42, 49 (2, 87.4 – 92.7 mm SL): “ Rio Capin ” (= Brazil, Pará, Rio Capim); “ purch. Stevens ” (i. e., collected by H. W. Bates); syntypes of Brycon brevicauda Günther, 1864. BMNH 1842.4. 20.13 (1, 232.0 mm SL): “ Rio Jocintins ” (= Rio Tocantins, Brazil); “ pres. by M. Gardiner ”; syntype of Brycon brevicauda Günther. MNHN 87746 - 748 (3, 110.0 – 120.9 mm SL): “ Orénoque ”, Chaffanjon. Syntypes of Brycon bicolor Pellegrin. ANSP 69619 (1, 149.2 mm SL): Brazil: “ Rio Parnahyba; Therezina; Piauhy ”; R. von Ihering, 1936. Holotype of Brycon matrinchao Fowler. Non types. French Guyana. MNHN 2000 - 4476 (1, 162.6 mm SL): Maroni River at Maripasoula, 3 ° 38 ’ 48 ’’ N, 54 ° 2 ’ 27 ’’ W; IRD-Cayenne, 15 June 1999. MNHN 1999 - 1451 (1, 102.0 mm SL): Maroni River, Saut Singatelet, c. 4 ° 20 ’ N, 54 ° 23 ’ W; P. Planquette, 15 Oct 1979. MNHN 2004 - 0345 (1, 99.1 mm SL); MNHN 2004 - 0243 (1, 70.9 mm SL): Saint Laurent du Maroni, 5 ° 28 ’ N, 54 ° 3 ’ W; O. Tostain, 2003. MNHN 1998 - 1822 (1, 257.9 mm SL): Saint Laurent du Maroni, 5 ° 28 ’ N, 54 ° 3 ’ W; P. - Y. Le Bail, no date. Suriname, Nickerie District: MZUSP 38255 (4, 1 cs, 116.9 – 131.6 mm SL): woodland stream about 0.5 km inland of Camp Mataway, Corantijn River basin, 4 ° 48 ' N, 57 ° 43 ' W; R. P. Vari, 21 Sept 1980. USNM 225975 (2, 129.1 – 152.0 mm SL): small creek entering Corantijn River, on east side approx. 300 m north of Amotopo boat landing, 3 ° 33 ’ N, 57 ° 40 ’ W; R. P. Vari et al., 19 Sept 1980. USNM 225625 (3, 134.7 – 141.2 mm SL): creek opposite logging camp, 2 and a half hours S of Matapi, approx. 2 km downstream Cow Falls, 4 ° 59 ’ N, 57 ° 38 ’ W; R. P. Vari et al., 11 Sept 1980. USNM 226161 (66, 77.1 – 108.1 mm SL): Corantijn River at km 180, side channel along Surinamese shore, 5 ° 8 ’ N, 57 ° 18 ’ W; R. P. Vari et al., Sept 1980. USNM 225624 (4, 127.6 – 136.6 mm SL): Mataway Creek approximately 8 km from its intersection with Corantjin River, 4 ° 47 ' N, 57 ° 45 ' W; R. P. Vari et al., 11 Sept 1980. USNM 226122 (9, 96.7 – 140.0 mm SL): Matapi Creek ca. 1 km from intersection with Corantijn River, 5 ° 00 ' N, 57 ° 16 ' W; R. P. Vari et al., 9 Sept 1980. AMNH 54849 (98.9 – 118.8 mm SL): stream near Camp Avanavaro, about 3 mi. downstream of Devis Falls (Kabalebo River drainage); R. P. Vari et al., 7 Dec 1979. AMNH 54976 (5, 103.4 – 114.2 mm SL): small stream just south of Tiger Falls on Corantijn River approx. km 405, c. 4 ° 0 ’ N, 58 ° 2 ’ W; R. P. Vari et al., 11 Dec. 1979. AMNH 54876 (7, 100.2 – 124.3 mm SL): Toeboeroe creek, trib. Corantjin River at km 220; R. P. Vari et al., 8 Dec 1979. AMNH 54764 (9, 97.4 – 113.7 mm SL): Corantijn drainage, Kapoeri Creek about 7 km from intersection with Corantjin River; R. P. Vari et al., 5 Dec 1979. AMNH 54904 (10, 90.0 – 105.9 mm SL): small inlets on sand bar opposite entrance of Matawai creek in Corantjin River; R. P. Vari et al., 8 Dec 1979. Guyana, Essequibo River basin. MCZ 30162 (1, 193.3 mm SL); FMNH 53352 (2, 196.5 – 222.0 mm SL): Tukeit, Potaro River, c. 5 ° 16 ’ N, 59 ° 23 ’ W; C. H. Eigenmann, 1908. FMNH 7472 (1, 188.8 mm SL): same locality; S. E. Shideler, 1908. AMNH 73000 (6, 82.5 – 124.9 mm SL): Mazaruni-Potaro District, Cuyuni River, just upstream of Caowry Creek, c. 6 ° 23 ’ N, 58 ° 43 ’ W; R. E. Schmidt, K. Schmidt & R. Pappantoniou, 18 Aug 1983. UMMZ 216328 (2, 102.1 – 107.0 mm SL): Essequibo River, Bartica, 6 ° 24 ’ N, 58 ° 37 ’ W; Carslon & Persand, 12 Sept 1971. UMMZ 216215 (3, 41.4 – 50.6 mm SL): small creek trib. Essequibo River, south of Bartica; F. Cichocki, 21 May 1972. UMMZ 216492 (2, 51.2 – 66.0 mm SL): Essequibo River, opposite Bartica; F. Cichocki & Douglas, 4 June 1972. UMMZ 216401 (1, 38.3 mm SL): Essequibo River, east bank up to Makouria Quarry; F. Cichocki et al., 18 May 1972. BMNH 1972.19.17: 1398 - 1411 (14, 82.7 – 125.6 mm SL): Essequibo River, Moraballi, c. 6 ° 12 ’ N, 58 ° 34 ’ W; R. Liley, 3 Nov 1959. FMNH 53350 (1, 89.7 mm SL): Essequibo River, Rockstone, 6 ° 1 ’ N, 58 ° 34 ’ W; C. H. Eigenmann, 1908. ANSP 177356 (2, 185.0 – 255.0 mm SL): Siparuni River, Levi Falls Creek; G. Watkins et al., 5 Dec 1997. ANSP 175740 (1, 250.0 mm SL): Essequibo River, approx. 3 hours above Kupurukari field station; W. G. Saul et al., 30 Jan 1997. ANSP 178559 (1, 248.0 mm SL): Burro Burro River: ca. 35 minutes upstream from Burro Burro campsite (Water Dog Rapids), 4 ° 10 ' 48 '' N, 58 ° 50 ' 46 '' W; G. G. Watkins et al., 22 Jan 1997. ANSP 175455 (1, 106.0 mm SL): Siparuni VIII- 2, blackwater creek tributary of Burro Burro River, upstream from Burro Burro campsite, 4 ° 42 ' 47 '' N, 58 ° 51 ' 46 '' W; G. G. Watkins et al., 21 Jan 1997. ANSP 175451 (1, 141.4 mm SL): Siparuni VIII- 2, clear water creek at campsite 3.1 miles from Kurupukari, field station on Kurupukari-Surema River Road (Tiger cr.), 4 ° 38 ' 00 '' N, 58 ° 42 ' 59 '' W; W. G. Saul et al., 5 Febr 1997. ANSP 176666 (1, 252.0 mm SL); ANSP 176667 (1, 165.3 mm SL): Burro Burro River, Water Dog Falls, 4 ° 10 ' 48 '' N, 58 ° 50 ' 46 '' W; E. McBirney et al., 17 – 20 Nov 1997. ANSP 176668 (1, 137.4 mm SL): Burro Burro River, Lunch Rock, 4 ° 41 ' 28 '' N, 58 ° 51 ' 8 '' W; C. Watson et al., 20 Nov 1997. ANSP 176667 (1, 165.3 mm SL): same locality; G. Watkins et al., 17 Nov 1997. ANSP 175456 (1, 134.4 mm SL): Siparuni VIII- 2, Paddle Rock lake, ca. 15 min downstream from Essequibo campsite, Essequibo River, 4 ° 43 ' 57 '' N, 58 ° 42 ' 53 '' W; W. G. Saul, 27 Jan 1997. ANSP 176669 (1, 135.2 mm SL): Essequibo River, black water creek 2 km downstream from Paddle Rock campsite, 4 ° 45 ' 40 '' N, 58 ° 43 ' 05 '' W; C. Watson et al., 24 Nov 1997. ANSP 175454 (1, 100.7 mm SL): Siparuni VIII- 2, Essequibo River, 180 yd. upstream from Essequibo campsite (Maipuri), 4 ° 45 ' 43 '' N, 58 ° 45 ' 52 '' W; D. Allicock, 27 Jan 1997. ANSP 176670 (2, 128.4 – 134.3 mm SL): Siparuni River, small side stream of Siparuni next to Tumble Down Falls, 4 ° 48 ' 39 '' N, 58 ° 51 ' 11 '' W; G. Watkins et al., 8 Dec 1997. ANSP 175457 (1, 121.3 mm SL): Siparuni VIII- 2, Essequibo River: sandbars in vicinity of Maipuri campsite, 4 ° 34 ' 17 '' N, 58 ° 35 ' 17 '' W; W. G. Saul et al., 2 Feb. 1997. ROM 64251 (2, 93.1 – 101.9 mm SL): Region 6 (Kurupukari), about 1.6 km downstream from Tambikabo inlet, 4 ° 48 ’ 40 ’’ N, 58 ° 49 ’ 20 ’’ W; E. Holm et al., 13 Oct 1990. ANSP 39752 (3, 86.3 – 109.1 mm SL): Rupununi River (no precise locality); J. Ogilvie, 1911. BMNH 1972.7.27: 52 - 76 (25, 79.5 – 255.0 mm SL): Rupununi District (no precise locality); R. H. Lowe McConnel, no date. MNHN A. 8614 (1, 244.3 mm SL, half skin); MNHN A. 9831 (1, 112.4 mm SL); MNHN 4388 (2, 110.0 – 115.8 mm SL); MNHN 101 (2, 205.3 – 218.3 mm SL): “ Fl. Essequibo, Guyane Anglaise ”; R. Schomburgk, no date. Venezuela, Río Orinoco basin, Depto. Amazonas: AMNH 93064 (1, 197.7 mm SL): Rio Mavaca, upriver from base camp, at tributary, c. 2 ° 31 ’ N, 65 ° 10 ’ W; C. J. Ferraris, A. Machado-Allison et al., 18 – 20 March 1989. AMNH 93065 (1, 202.8 mm SL): Río Mavaca, at base camp, c. 2 ° 31 ’ N, 65 ° 10 ’ W; C. J. Ferraris & J. Daly, 17 March 1989. ANSP 159731 (1, 114.0 mm SL): Río Sipapo, above Pendare, 4 ° 51 ’ N, 67 ° 44 ’ W; B. Chernoff et al., 12 Nov 1985. ANSP 159734 (1, 91.5 mm SL): Caño entering Río Sipapo at raudal del Caldero, c. 3 km above confluence with Río Orinoco, 5 ° 4 ’ N, 67 ° 46 ’ W; B. Chernoff et al., 14 Nov 1985. ANSP 159732 (1, 110.1 mm SL): morichal 26.9 km from Puerto Ayacucho, along Puerto Ayacucho- Caicara highway; B. Chernoff et al., 15 Nov 1985. ANSP 159733 (1, 127.2 mm SL): Río Cataniapo at bridge on Puerto Ayacucho-Samariapo carretera, 3 km S of Puerto Ayacucho, 5 ° 32 ' N 67 ° 31 ' W; B. Chernoff et al., 11 Nov 1985. FMNH 85689 (4, 2 cs, 92.4 – 147.7 mm SL): 50 km towards Puerto Ayacucho from Puerto Novo; J. Thomerson et al., 14 Jan 1975. USNM 270181 (3, 97.3 – 120.0 mm SL): Laguna Provincial, approx. 20 km north of Puerto Ayacucho, 5 ° 50 ’ N, 67 ° 30 ’ W; R. P. Vari et al., 1 Dec 1984. INHS 61580 (1, 118.3 mm SL): Caño Agua Linda (trib. Río Orinoco), 5 ° 50 ' 24 '' N 67 ° 27 ' 10 '' W; L. M. Page et al., 22 Jan 1992. ANSP 159730 (2, 83.9 – 97.7 mm SL): Caño crossing Puerto Ayacucho-El Burro hwy., c. 7 km from intersection of Puerto Ayacucho-Caicara hwy., 5 ° 56 ’ N, 67 ° 21 ’ W; B. Chernoff et al., 14 Nov 1985. ANSP 159721 (1, 109.3 mm SL): Río Orinoco at El Burro, 6 ° 12 ’ N, 67 ° 26 ’ W; B. Chernoff et al., 26 Nov 1985. BMNH uncat. (1, 86.5 mm SL): Río Orinoco, Puerto Ayacucho, 5 ° 40 ’ N, 67 ° 38 ’ W; Delmastro, July 1980. UMMZ 240018 (3, 94.8 – 136.0 mm SL): mouth of Río Guaviare, opposite San Fernando de Atabapo, 4 ° 2 ’ N, 67 ° 43 ’ W; J. S. Albert et al., 2 Jan 1994. UMMZ 239924 (1, 93.7 mm SL): Río Orinoco, 5 km upstream from San Fernando de Atabapo, c. 4 ° 2 ’ N, 67 ° 40 ’ W; J. S. Albert et al., 27 Dec 1993. ANSP 161212 (8, 106.4 – 149.8 mm SL): Río Ventuari ca. 12 km from its confluence with Río Orinoco, 4 ° 4 ’ N, 66 ° 56 ’ W; B. Chernoff et al., 25 March 1987. FMNH 104026 (1, 100.4 mm SL); FMNH 104027 (1, 92.6 mm SL); FMNH 104028 (1, 117.5 mm SL): Río Ventuari, c. 12 km above mouth in Río Orinoco, Laguna Pavón, 4 ° 4 ’ N, 66 ° 56 ’ W; B. Chernoff et al., 24 Jan 1991. FMNH 161210 (1, 157.1 mm SL): Río Iguapo (trib. Río Orinoco), c. 1 hr. above its mouth, 3 ° 7 ’ N, 65 ° 28 ’ W; H. Lopez et al., 13 March 1987. CAS-SU 52 635 (1, 102.8 mm SL): Rio Orinoco, Orinoco bifurcation, Tamatama beach, c. 3 ° 9 ’ N, 65 ° 51 ’ W; C. Ternetz, 14 March 1925. CAS-SU 64370 (1, 230 mm SL): Rio Orinoco bifurcation, Cano Tamatama into Rio Orinoco, c. 3 ° 9 ’ N, 65 ° 51 ’ W; C. Ternetz, 17 March 1925. Estado Bolívar: ANSP 159736 (1, 100.7 mm SL): Río Caura at Puerto Las Majadas, 7 ° 38 ’ N, 64 ° 50 ’ W; L. Aguana, 23 Nov 1985. MZUSP 96418 (1, 54.2 mm SL): Caicara del Orinoco, Río Orinoco, Puerto Cedeño, 7 ° 39 ’ 6 ’’ N, 66 ° 10 ’ 34 ’’ W; M. C. C. de Pinna & P. Hernadez, 20 July 2004. LBP 2187 (2, 98.4 – 147.6 mm SL): Caicara del Orinoco, Laguna de Castilleros, 7 ° 30 ’ 51 ’’ W, 66 ° 9 ’ 20 ’’ W; A. Granado, 17 Jul 2004. LBP 3027 (4, 101.2 – 172.0 mm SL): Caicara del Orinoco, Río Orinoco, 7 ° 38 ’ 11 ’’ N, 66 ° 19 ’ 4 ’’ W; C. Oliveira & A. Granado, 2 Oct 2005. CAS-SU 68916 (1, 107.8 mm SL); CAS-SU 54649 (1, 101.4 mm SL); CAS-SU 56 784 (1, 103.3 mm SL); CAS-SU 54566 (1, 104.5 mm SL): Caño de Quiribana, near Caicara, c. 7 ° 36 ’ N, 66 ° 11 ’ W; C. Ternetz, April – May 1925. UF 80419 (1, 112.1 mm SL); UF 80474 (1, 125.3 mm SL): bridge on Río Chaviripa on Caicara-San Fernando de Atabapo (Puerto Ayacucho) road, c. 7 ° 0 ’ N, 66 ° 31 ’ W; D. C. Taphorn et al., 16 Apr 1984. ANSP 159729 (1, 102.9 mm SL): river and flooded area 15 km N of Maniapure on Caicara-Puerto Ayacucho hwy. (Río Chaviripa?), c. 7 ° 0 ’ N, 66 ° 31 ’ W; B. Chernoff et al., 16 Nov 1985. MZUSP 96499 (1, 69.2 mm SL): Cedeño, Río Parguaza, Puente Parhueña village, 5 ° 53 ’ 30 ’’ N, 67 ° 24 ’ 14 ’’ W; M. C. C. de Pinna & C. Oliveira, 19 Jul 2004. Estado Apure: MZUSP 62447 (2, 104.6 – 124.4 mm SL): Apure, laguna near Caño La Pica; F. Provenzano et al., 14 May 1989. INHS 61479 (3, 84.6 – 113.5 mm SL): Apure, Caño San Miguel (trib. Río Cínaruco), 6 ° 34 ’ 24 ’’ N 67 ° 17 ’ 32 ’’ W; L. M. Page et al., 20 Jan 1992. INHS 61402 (1, 115.1 mm SL): Laguna Larga (Río Cínaruco), 6 ° 33 ’ 19 ’’ N 67 ° 24 ’ 49 ’’ W; L. M. Page et al., 19 Jan 1992. FMNH 69901 (1, 89.3 mm SL); FMNH 69902 (8, 94.1 – 169.4 mm SL): Río Cinaruco, c. 6 ° 33 ’ N, 67 ° 18 ’ W; W. P. Braker & Zunwalt, Feb 1967. Venezuela, Depto. Amazonas, Rio Negro basin: ANSP 161211 (1, 102.4 mm SL): Río Pamoni, lagoon ca. 0.5 km from confluence of Río Casiquiare, 2 ° 50 ’ N, 65 ° 53 ’ W; B. Chernoff et al., 18 March 1987. ANSP 162802 (1, 272.0 mm SL): mouth of Río Pamoni at Río Casiquiare, 2 ° 49 ’ N, 65 ° 55 ’ W; W. G. Saul et al., 18 March 1987. CAS-SU 56148 (1, 109.5 mm SL); CAS-SU 54738 (1, 127.7 mm SL); CAS-SU 54652 (2, not measured); CAS-SU 54652 (2, 107.9 – 117.2 mm SL): Rio Casiquiare, Laje Curare (not located); C. Ternetz, 2 March 1925. CAS-SU 54653 (2, 113 – 124.6 mm SL): Rio Casiquiare, Punta de la Isla Guachancho; C. Ternetz, 26 Febr 1925. USNM 270180 (1, 105.4 mm SL): San Carlos de Río Negro, Río Negro above town landing, 1 ° 55 ’ N, 67 ° 3 ’ W; A. Machado et al., 3 Dec 1984. USNM 270182 (5, 80.8 – 99.8 mm SL): Río Negro about 0.5 hour above San Carlos de Río Negro, 1 ° 58 ’ N, 67 ° 4 ’ W; R. P. Vari et al., 4 Dec 1984. CAS-SU 54651 (1, 109.8 mm SL); CAS-SU 69885 (1, 106.0 mm SL): Rio Negro, sandy point n. of “ Amanadona ” (= Santa Rosa de Amanadona, 1 ° 28 ’ 30 ’’ N, 66 ° 54 ’ 30 ’’ W); C. Ternetz, 21 Febr 1925. Colombia, Río Orinoco basin: NRM 26206 (1, 70.5 mm SL): Prov. Guainía, Río Guaviare drainage, Caño Carbón, Cuayare, 3 ° 56 ' N 67 ° 50 ' W; T. Hongslo, 28 June 1976. Brazil, rio Capim basin: MZUSP 17894 (1, 115.1 mm SL): Pará, São Domingos do Capim, igarapé Pirajauara, trib. rio Capim, c. 1 ° 44 ’ S, 47 ° 47 ’ W; EPA, 12 Aug 1970. MZUSP 17957 (5, 79.2 – 92.4 mm SL): Pará, Badajós, lago Maria Preta, rio Capim, c. 2 ° 30 ’ S, 47 ° 40 ’ W; EPA, 18 Aug 1970. Rio Tocantins basin. Pará: MZUSP 18051 (9, 143.8 – 160.1 mm SL): Paraná Samuuma, mouth of rio Tocantins, c. 1 ° 56 ’ S, 49 ° 12 ’ W; EPA, 4 Sept 1970. MZUSP 18068 (15, 100.4 – 127.6 mm SL): Mocajuba, igarapé Oxipucu, 2 ° 34 ’ S, 49 ° 31 ’ W; EPA, 8 Sept 1970. MZUSP 18087 (30, 1 cs, 98.1 – 113.9 mm SL): rio Tocantins, between Mocajuba and Baião, c. 2 ° 39 ’ S, 49 ° 37 ’ W; EPA, 9 Sept 1970. INPA 16424 (1, 191.9 mm SL); INPA 16368 (1, 119.7 mm SL); INPA 16386 (1, 168.0 mm SL); INPA 16382 (1, 146.4 mm SL): rio Tocantins, Acarí Pucu, 2 ° 42 ’ S, 49 ° 43 ’ W; Eq. Ictiologia / INPA, 1981 – 1982. MZUSP 18089 (5, 106.4 – 112.0 mm SL): Baião, rio Tocantins, 2 ° 48 ’ S, 49 ° 41 ’ W; EPA, 9 – 10 Sept 1970. MZUSP 18077 (21, 103.4 – 135.3 mm SL): Baião, igarapé do Limão, rio Tocantins; EPA, 9 Sept 1970. MZUSP 18195 (3, 100.3 – 105.0 mm SL): igarapé dos Cinco, km 5 on road Tucuruí-Mato Grosso; EPA, 22 Sept 1970. MZUSP 18200 (1,111.5 mm SL): igarapé Urubu, near posto Trocará, rio Tocantins; EPA, 24 Sept 1970. INPA 16392 (2, 206.7 – 210.9 mm SL); INPA 16373 (2, 141.4 – 146.7 mm SL): Icangui, rio Tocantins, 3 ° 27 ’ S, 49 ° 36 ’ W; Eq. Ictiologia / INPA, 1980 – 1985. MZUSP 18115 (4, 112.9 – 196.6 mm SL): marginal lagoons of rio Tocantins, near Tucuruí; EPA, 14 Sept 1970. MZUSP 61991 (1, 109.9 mm SL): Tucuruí, rio Tocantins, downstream Tucuruí dam, 3 ° 42 ' S 49 ° 40 ' W; F. C. T. Lima, 17 May 2000. INPA 16448 (1, 192.6 mm SL): rio Tocantins, Tucuruí fish market; Eq. Ictiologia / INPA, 13 March 1982 .. INPA 16452 (5, 43.3 – 73.5 mm SL): rio Tocantins, Tucuruí reservoir, c. 4 ° 2 ’ S, 49 ° 40 ’ W; Martinho, 13 March 1986. INPA 16431 (3, 185.6 – 199.1 mm SL): rio Tocantins, Breu Branco, c. 4 ° 11 ’ S, 49 ° 37 ’ W; Eq. Ictiologia / INPA, 13 May 1981. INPA 16387 (1, 95.3 mm SL): rio Tocantins, igarapé Pucuruizinho; Eq. Ictiologia / INPA, 9 July 1982. MZUSP 18145 (1, 84.3 mm SL); MZUSP 18134 (3, 89.3 – 98.9 mm SL): pool facing Jatobal, rio Tocantins, 4 ° 32 ’ S, 49 ° 32 ’ W; EPA, 17 – 18 Sept 1970. MZUSP 18163 (10, 88.9 – 111.1 mm SL): lagoon near canal do Capitariquara, near Jatobal, rio Tocantins; EPA, 18 Sept 1970. INPA 16362 (1, 109.0 mm SL);: Itupiranga, rio Tocantins, 5 ° 8 ’ S, 49 ° 19 ’ W; Eq. Ictiologia / INPA, 1980 – 1981. INPA 16381 (1, 84.6 mm SL): Itupiranga, rio Tocantins, Lago Gracílio; Eq. Ictiologia / INPA, 1 July 1982. MZUSP 26924 (1, 96.8 mm SL): Marabá, rio Tocantins, 5 ° 9 ’ S, 49 ° 6 ’ W; N. J. H. Smith, April 1979. MZUSP 19260 (1, 193.8 mm SL): São João do Araguaia, at the confluence of rio Araguaia and rio Tocantins, 5 ° 22 ’ S, 48 ° 43 ’ W; SUDEPE, 1977 – 1978. MZUSP 31503 (1, 218.9 mm SL): Parauapebas, rio Itacaiúnas, igarapé Boa Vista; M. Goulding, Nov 1983. MZUSP 56946 (2, 199.2 – 211.6 mm SL); MZUSP 56943 (3, 194.3 – 212.9 mm SL): Parauapebas, rio Itacaiúnas, Caldeirão, Serra dos Carajás, c. 5 ° 40 ’ S, 50 ° 16 ’ W; M. Goulding, April – July 1983. MZUSP 59100 (5, 157.5 – 164.1 mm SL): Parauapebas, Serra dos Carajás, rio Itacaiúnas, Cachoeira Carreira Comprida; M. Goulding, 14 Oct 1983. Mato Grosso: LBP 4005 (4, 113.3 – 125.0 mm SL): São Félix do Araguaia, Lago Morto, 11 ° 40 ’ 0 ’’ S, 50 ° 51 ’ 00 ’’ W; R. Devidé & C. Martins, 24 Oct 2006. MZUSP 17023 (1, 111.8 mm SL); MZUSP 17024 (1, 113.7 mm SL): Santa Teresinha, rio Araguaia, 10 ° 28 ’ S, 50 ° 30 ’ W; H. A. Britski, Oct 1964. MZUSP 54444 (2, 127.1 – 172.2 mm SL): rio Cristalino, c. 12 ° 43 ’ S, 50 ° 42 ’ W; R. A. Silvano, 3 Oct 1997. MZUSP 16448 (24, 121.1 – 209.1 mm SL): rio São Domingos, trib. rio das Mortes, São Domingos village, 13 ° 33 ’ S, 51 ° 25 ’ W; Exp. Inst. Butantã / Depto. de Zoologia, Sept – Oct 1949. MZUSP 3845 (15, 130.1 – 215.7 mm SL): same locality; A. Hoge, 1950. MZUSP 52332 (1, 190.9 mm SL): Cocalinho, rio Araguaia, 14 ° 24 ’ S, 50 ° 59 ’ W; R. S. A. Matias, July 1997. LBP 12816 (1, 97.9 mm SL): Cocalinho, rio Araguaia, 13 ° 18 ’ 37 ’’ S, 50 ° 36 ’ 48 ’’ W; R. Devidé et al., 29 Sept 2009. MZUSP 62537 (3, 172.0 – 197.7 mm SL): Cocalinho, rio Araguaia, near ilha do Biratã, c. 14 ° 30 ’ S, 50 ° 59 ’ W; W. Severi et al., 25 – 26 July 1997. MZUSP 18628 (2, 184.6 – 214.7 mm SL): Cocalinho, Lago Dumbá, rio Araguaia, c. 14 ° 30 ’ S, 50 ° 59 ’ W; EMGOPA, 1976. Maranhão: CAS 68839 (1, 123.2 mm SL): rio Tocantins below Imperatriz, 5 ° 30 ’ S, 47 ° 31 ’ W; C. Ternetz, 16 Apr 1924. Tocantins: INPA 20064 (1, 126.3 mm SL): Caseara, rio Araguaia, lago das Ariranhas, P. E. Cantão, 9 o 43 ’ 47 ” S, 50 o 9 ’ 14 ” W; Eq. Ictiologia INPA, 20 May 2000. UNT 448 (1, 155.4 mm SL): Dueré, rio Formoso, c. 11 ° 23 ’ S, 49 ° 41 ’ W; NEAMB, 23 Aug 1997. UNT 1971 (1, 107.3 mm SL): Tocantinópolis, ribeirão Matrinchã, c. 6 ° 19 ’ S, 47 ° 26 ’ W; NEAMB, 26 June 2000. CAS 68824 (1, 214 mm SL); CAS 68825 (4, 181.5 – 192.3 mm SL): Ribeirão Presídio, trib. Rio Tocantins, below Porto Nacional; C. Ternetz, 13 Feb 1924. UNT 3457 (1, 129.7 mm SL): rio Tocantins, Porto Nacional, 10 ° 42 ’ S, 48 ° 25 ’ W; NEAMB, 21 Feb 2002. UNT 3225 (1, 96.6 mm SL): Brejinho de Nazaré, riacho Sussuarana, c. 11 ° 3 ’ S, 48 ° 35 ’ W; NEAMB, 15 Feb 2002. UNT 2013 (1, 131.3 mm SL): Peixe, rio Santa Tereza (trib. rio Tocantins), 11 ° 47 ’ S, 48 ° 38 ’ W; NEAMB, 12 April 2000. UNT 451 (1, 193.8 mm SL); UNT 452 (1, 221.4 mm SL): Paranã, rio Paranã, 12 ° 37 ’ S, 47 ° 53 ’ W; NEAMB, 24 – 26 March 1998. UNT 1970 (1, 106.4 mm SL): Paranã, rio Maranhão; NEAMB, 21 Apr 1998. Goiás: MCP 17213 (1, 1, 190.2 mm SL): Luís Alves, rio Araguaia, marginal lagoons, 13 ° 14 ' S, 50 ° 35 ' W; F. L. T. Garro, 21 April 1994. MZUSP 89614 (1, 111.7 mm SL): Luís Alves, rio Verde (trib. rio Araguaia), 13 ° 9 ’ 28 ’’ S, 50 ° 30 ’ 22 ’’ W; A. Akama, 10 Apr 2004. MZUSP 52391 (1, 200.9 mm SL): rio Araguaia, Bandeirantes, 13 ° 41 ’ S, 50 ° 48 ’ W; R. S. A. Matias, July 1997. MZUSP 48113 (4, 197.7 – 224.6 mm SL): Aragarças, rio Araguaia, c. 15 ° 54 ’ S, 52 ° 15 ’ W; W. P. Margarido, 6 July 1994. MZUSP 70418 (1, 179.5 mm SL): Serra da Mesa dam, rio Palmeirinha, 14 ° 03 ' 57 ' ’ S, 48 ° 29 ' 37 ' ’ W; D. F. Moraes & D. A. Halboth, 8 June 1997. MZUSP 70415 (1, 243.7 mm SL): rio Maranhão, old bridge between Uruaçu and Niquelândia, 14 ° 31 ' 27 ' ’ S, 49 ° 2 ' 33 ' ’ W; D. F. Moraes et al., 7 April 1998. MZUSP 70416 (1, 85.4 mm SL): rio Maranhão, between the mouth of rio Bagagem and rio Tocantizinho, 13 ° 56 ' S, 48 ° 17 ' W; D. F. Moraes & D. A. Halboth, 6 Dec 1996. MZUSP 70417 (2, 109.9 – 245.6 mm SL): Serra da Mesa dam, córrego Boa Nova, 13 ° 47 ' 00 ' ’ S, 48 ° 25 ' 01 ' ’ W; D. F. Moraes, 6 Feb 1997. Rio Xingu basin. Pará: MZUSP 29922 (3, 59.2 – 97.9 mm SL); MZUSP 56945 (151, 117.2 – 193.5 mm SL); MZUSP 31448 (1, 153.5 mm SL); MZUSP 56811 (1, 155.7 mm SL); MZUSP 56947 (1, 194.3 mm SL); MZUSP 30755 (2, 82.6 – 93.8 mm SL); MZUSP 58985 (3, 97.1 – 100.9 mm SL): Belo Monte, rio Xingu, 3 ° 7 ' S 51 ° 42 ' W; M. Goulding, July – Aug 1983. MZUSP 29923 (3, 89.5 – 129.8 mm SL); MZUSP 58986 (4, 93.6 – 107.8 mm SL): Belo Monte, rio Xingu (rocky pool), 3 ° 7 ' S 51 ° 42 ' W; M. Goulding, 26 Sept 1983. MZUSP 29928 (1, 103.4 mm SL): igarapé do Santo Antônio, near Belo Monte, rio Xingu; M. Goulding, 27 Sept 1983. INPA 15747 (4, 121.5 – 130.4 mm SL); INPA 4225 (1, 86.4 mm SL); INPA 15745 (1, 166.5 mm SL): rio Xingu, Altamirim, ilha do Sr. Izaltino, 3 ° 16 ’ 21 ’’ S, 52 ° 12 ’ 7 ’’ W; J. Zuanon, Oct 1996 – Aug 1997. INPA 15748 (1, 149.2 mm SL): rio Xingu, Souzel, ilha da Bela Vista, 3 ° 24 ’ 22 ’ S, 51 ° 43 ’ 3 ’’ W; J. Zuanon et al., 27 – 28 Sept 1996. ZUEC 4487 (1, 98.9 mm SL): Altamira, rio Xingu, Buraco do Inferno; J. Zuanon, 7 Sept 1997. INPA 4226 (4, 89.6 – 95.9 mm SL): rio Xingu, Ilha de Babaguara; L. Rapp Py-Daniel & J. Zuanon, 5 Oct 1990. MZUSP 105722 (1, 170.7 mm SL): Altamira, rio Xingu, Caitucá, 3 ° 33 ’ 48 ’’ S, 51 ° 51 ’ 49 ’’ W; Eq. Ictiologia UFPA, 14 Sept 2001. MNHN 1998 - 1197 (3, 97.1 – 181.1 mm SL): Altamira, rio Xingu, Caitucá, 3 ° 33 ’ 48 ’’ S, 51 ° 51 ’ 49 ’’ W; M. Jégu, Oct 1992. MZUSP 106211 (1, 159.0 mm SL): Altamira, rio Xingu, Paquiçamba, 3 ° 33 ’ 44 ’ S, 51 ° 52 ’ 37 ’’ W; Eq. Ictiologia UFPA, 5 – 6 Nov 2000. MZUSP 36792 (3, 106.9 – 162.2 mm SL): rio Xingu, Cachoeira do Espelho, 3 ° 39 ’ 2 ’’ S, 52 ° 22 ’ 46 ’’ W; P. E. Vanzolini, 23 – 26 Oct 1986. Mato Grosso: MZUSP 94946 (2, 195.0 – 325.0 mm SL): Campinápolis, Rio Sucuri, trib. rio Culuene, 13 ° 55 ’ 40 ’’ S, 53 ° 17 ’ 10 ’’ W; A. Akama & J. L. Birindelli, 15 – 22 Jan 2006. MZUSP 91924 (1, 318.0 mm SL): Campinápolis, rio Culuene, below future PCH Paranatinga II, 13 ° 49 ’ S, 53 ° 15 ’ W; J. L. Birindelli et al., 21 Aug 2006. MZUSP 94962 (3, 247.0 – 347.0 mm SL): same locality; F. C. T Lima et al., 19 – 20 May 2007. MZUSP 94897 (21, 4 skel., 278.0 – 355.0 mm SL): same locality; L. M. Sousa et al., July 2007. MZUSP 98119 (13, 311.0 – 380.0 mm SL): same locality; F. C. T. Lima et al., Oct 2007. MZUSP 91857 (1, c. 300 mm SL): Córrego do Lício, trib. rio Culuene, below PCH Paranatinga II, 13 ° 50 ’ 22 ’’ S, 53 ° 14 ’ 59 ’’ W; J. L. Birindelli et al., 21 Aug 2006. MZUSP 98118 (2, 297.0 – 336.0 mm SL): Campinápolis, rio Culuene, rio Corgão, at its mouth, 13 ° 48 ’ 7 ’’ S, 53 ° 15 ’ 0 ’’ W; F. C. T. Lima et al., 4 – 14 Oct 2007. MZUSP 98131 (2, 309.0 – 401.0 mm SL): Campinápolis, rio Culuene, Cachoeira do Adelino, 13 ° 47 ’ 50 ’’ S, 53 ° 14 ’ 46 ’’ W; F. C. T. Lima et al., 2 – 14 Oct 2007. MZUSP 94377 (3, 112.6 – 187.3 mm SL): Canarana, lake at fazenda Miriam, rio Culuene, 13 ° 25 ’ 48 ’’ S, 53 ° 2 ’ 24 ’’ W; F. C. T. Lima et al., 23 May 2007. MZUSP 56809 (1, 233.6 mm SL): Gaúcha do Norte, rio Xingu, confluence of rio Culuene and rio Sete de Setembro, 12 ° 55 ’ 28 ’’ S, 52 ° 49 ’ 34 ’’ W; M. Goulding et al., 23 Aug 1984. MZUSP 103199 (1, 376.0 mm SL): Nova Ubiratã, rio Von den Steinen, fazenda A. R. S., 13 ° 5 ’ 35 ’’ S, 54 ° 49 ’ 8 ’’ W; F. A. Machado et al., 21 Jan 2006. Rio Tapajós basin. Pará: MZUSP 18255 (8, 188.4 – 210.1 mm SL); MZUSP 21988 (2, 92.7 – 117.7 mm SL): São Luís, rio Tapajós, 4 ° 27 ' S 56 ° 15 ' W; EPA, 4 – 8 Nov 1970. MZUSP 56955 (5, 159.5 – 182.3 mm SL); MZUSP 30759 (1, 114.7 mm SL): rio Tapajós, between Itaituba and São Luís, c. 4 ° 23 ’ S, 56 ° 5 ’ W; M. Goulding, Sept – Oct 1983. MZUSP 58095 (8, 103.2 – 146.0 mm SL): Itaituba, rio Tapajós above Itaituba (rapids); M. Goulding, 22 Oct 1983. MZUSP 25585 (1, 175.7 mm SL): rio Tapajós, between National Park headquarters and Ramal Saita, km 67, Parque Nacional da Amazônia, 4 ° 33 ’ S, 56 ° 19 ’ W; J. C. Oliveira, 4 – 6 Jan 1979. MZUSP 22026 (8, 99.3 – 122.9 mm SL): lago da Santa Clara, Monte Cristo, rio Tapajós, c. 4 ° 1 ’ S, 55 ° 39 ’ W; EPA, 6 Dec 1970. MZUSP 25362 (3, 88.6 – 101.0 mm SL): igarapé Mambuaí, bridge at BR- 230, Amazônia National Park, rio Tapajós; J. C. Oliveira, 15 – 31 Jul 1979. MZUSP 25452 (1, 94.2 mm SL): rio Tapajós, Porto Flexal, below Amazônia National Park headquarters, c. 4 ° 37 ’ S, 56 ° 18 ’ W; J. C. Oliveira, 15 – 31 July 1979. MZUSP 25436 (1, 94.2 mm SL); MZUSP 25437 (1, 84.4 mm SL): Ilha Grande, rio Tapajós, above Amazônia National Park headquarters; J. C. Oliveira, 15 – 31 July 1979. MZUSP 18251 (2, 143.3 – 206.0 mm SL): rio Tapajós, Barreirinha; EPA, Nov 1970. MZUSP 18287 (5, 123.7 – 137.1 mm SL): lake facing Monte Cristo, rio Tapajós; EPA, 8 Dec 1970. MZUSP 21876 (1, 95.5 mm SL): Maloquinha, near Itaituba, rio Tapajós; EPA, 11 – 13 Nov 1970. MZUSP 22081 (16, 115.4 – 135.3 mm SL): ilha da Barrerinha, rio Tapajós, near São Luís; EPA, 21 Nov 1970. MZUSP 25504 (1, 139.9 mm SL): Furo do Sandro, rio Tapajós, above Pimental, Amazônia National Park; J. C. Oliveira, 6 Jan 1979. MZUSP 25619 (2, 126.3 – 155.0 mm SL): rio Tapajós, between Ramal Saita and igarapé da Boa Vista, Amazônia National Park; J. C. Oliveira, 16 Jan 1979. MZUSP 29926 (8, 1 cs, 91.0 – 103.4 mm SL): rio Tapajós, Pederneiras, below Itaituba; M. Goulding, 14 Oct 1983. INPA 16410 (2, 146.5 – 159.7 mm SL); INPA 7058 (1, 131.5 mm SL): rio Tapajós, Pimental, 4 ° 34 ’ 29 ’’ S, 56 ° 15 ’ 44 ’’ W; L. H. Rapp Py-Daniel & J. Zuanon, 22 – 24 Oct 1991. MZUSP 92804 (1, 134.8 mm SL): Itaituba, rio Tapajós, 4 ° 16 ’ 14 ’’ S, 55 ° 58 ’ 34 ’’ W; J. L. Birindelli & L. M. Sousa, 7 Nov 2006. LBP 12871 (3, 147.4 – 225.0 mm SL): Itaituba, rio Tapajós, 4 ° 33 ’ 10 ’’ S, 56 ° 18 ’ 00 ’’ W; R. Britzke & CEPTA team, 24 Sept 2011. INPA 7292 (1, 132.3 mm SL): rio Tapajós (no specific locality); L. H. Rapp Py-Daniel & J. Zuanon, Oct 1991. INPA 6969 (18, 85.8 – 129.6 mm SL): rio Tapajós, Pimental (rocky pool); L. H. Rapp Py-Daniel et al., 23 Oct 1991. INPA 6793 (4, 93.7 – 144.0 mm SL); INPA 6636 (2, 107.7 – 132.3 mm SL): rio Jamanxim, trib. rio Tapajós, Ilha Terra Preta, 4 ° 47 ’ S, 56 ° 24 ’ W; L. H. Rapp Py-Daniel & J. Zuanon, 19 – 20 Oct 1991. MZUSP 99926 (3, 85.0 – 102.9 mm SL): trib. rio Teles Pires, below Sete Quedas rapids, 9 ° 18 ’ 27 ’’ S, 56 ° 47 ’ 38 ’’ W; L. M. Sousa & A. L. Netto-Ferreira, 9 June 2008. MZUSP 99874 (1, 110.3 mm SL): rio Teles Pires, below Sete Quedas rapids, 9 ° 18 ’ 42 ’’ S, 56 ° 46 ’ 47 ’’ W; L. M. Sousa & A. L. Netto-Ferreira, 9 June 2008. MZUSP 99564 (1, 42.5 mm SL); MZUSP 99512 (3, 191.7 – 192.1 mm SL): rio Teles Pires, near Sete Quedas rapids, 9 ° 20 ’ 20 ’’ S, 56 ° 46 ’ 30 ’’ W; R. Hilário, 23 – 25 March 2008. Mato Grosso: MZUSP 100055 (3, 72.2 – 101.1 mm SL): Paranaíta, rio Teles Pires, above Sete Quedas, 9 ° 23 ’ 53 ’’ S, 56 ° 34 ’ 37 ’’ W; L. M. Sousa & A. L. Netto-Ferreira, 16 June 2008. MZUSP 99996 (2, 94.6 – 110.9 mm SL): Paranaíta, rio Teles Pires, above Sete Quedas, 9 ° 25 ’ 2 ’’ S, 56 ° 33 ’ 1 ’’ W; L. M. Sousa & A. L. Netto-Ferreira, 16 June 2008. MZUSP 99574 (1, 44.3 mm SL): Paranaíta, rio Teles Pires, 9 ° 25 ’ 5 ’’ S, 56 ° 32 ’ 21 ’’ W; R. Hilário, 2 April 2008. MZUSP 95620 (14, 97.0 – 197.0 mm SL): Paranaíta, rio Teles Pires, near ferry at MT- 416 road, 9 ° 27 ’ 7 ’’ S, 56 ° 30 ’ 46 ’’ W; L. M. Sousa & A. L. Netto-Ferreira, 27 Sept 2007. MZUSP 106177 (8, 250.0 – 405.0 mm SL): Alta Floresta, rio Cristalino, P. E. do Cristalino, c. 9 ° 27 ’ S, 55 ° 50 ’ W; S. A. A. Silva, 2004 – 2006. MZUSP 100017 (1, 120.0 mm SL): Novo Mundo, rio Nhandu (trib. rio Teles Pires), below Nhandu dam, c. 9 ° 53 ’ S, 55 ° 22 ’ W; K. de Silimon, 19 Nov 2006. MZUSP 62537 (1, 106.7 mm SL): Alta Floresta, rio Teles Pires, pesqueiro do Dentinho, c. 10 ° 15 ’ S, 55 ° 49 ’ W; F. A. Machado et al., 29 – 30 July 1997. MZUSP 62508 (2, 150.5 – 174.8 mm SL): Alta Floresta, rio Teles Pires, Porto do Alcindo (1 km below mouth of rio Peixoto de Azevedo), 9 ° 59 ' 25 ' ’ S, 55 ° 33 ' 48 ' ’ W; F. A. Machado et al., 29 – 30 July 1997. MZUSP 96822 (2, 210.0 – 220.0 mm SL): Peixoto de Azevedo, Cachoeira da Neblina, trib. rio Peixoto de Azevedo, 10 ° 23 ’ 10 ’’ S, 54 ° 18 ’ 22 ’’ W; J. L. Birindelli et al., 18 Oct 2007. MZUSP 95866 (2, 157.5 – 159.2 mm SL): Itaúba, rio Teles Pires, 10 ° 58 ’ 30 ’’ S, 55 ° 44 ’ 3 ’’ W; J. L. Birindelli & P. H. Carvalho, 1 Oct 2007. MZUSP 99135 (1, 290.0 mm SL): Itaúba, rio Renato (trib. rio Teles Pires), 11 ° 4 ’ 24 ’’ S, 55 ° 14 ’ 35 ’’ W; P. Hollanda-Carvalho & S. Lima, 26 Feb 2008. MZUSP 99385 (1, 392.0 mm SL): Itaúba, rio Teles Pires, below mouth of rio Renato, 11 ° 3 ’ 44 ’’ S, 55 ° 19 ’ 8 ’’ W; P. Hollanda-Carvalho & S. Lima, 19 Feb 2008. MZUSP 62559 (1, 417.9 mm SL): Sinop, rio Teles Pires, Ilha do Peixeirinho, 11 ° 34 ’ 55 ’’ S, 55 ° 38 ’ 53 ’’ W: F. A. Machado et al., 1 – 2 Aug 1997. ZUEC 9190 (1, 345.0 mm SL): Sinop, rio Teles Pires, 11 ° 35 ’ 17 ’’ S, 55 ° 39 ’ 36 ’’ W; L. Matos & L. N. Carvalho, 19 Nov 2014. MZUSP 82015 (1, 409.0 mm SL): Lucas do Rio Verde, rio Verde (trib. rio Teles Pires), 13 ° 3 ’ S, 55 ° 54 ’ W; K. de Silimon, 16 March 2001. MZUSP 56785 (2, 199.5 – 233.6 mm SL): Porto dos Gaúchos, rio Arinos, 11 ° 32 ’ S, 57 ° 25 ’ W; M. Goulding, 19 Aug 1984. MZUSP 92633 (1, 279.0 mm SL): Sapezal, rio Juruena, below future PCH Telegráfica, 12 ° 41 ’ 22 ’’ S, 58 ° 56 ’ 47 ’’ W; K. de Silimon et al., 15 Sept 2006. MZUSP 95506 (1, 271.0 mm SL): Sapezal, rio Papagaio at mouth of rio Buriti, road Sapezal / Brasnorte, 12 ° 47 ’ 6 ’’ S, 58 ° 23 ’ 5 ’’ W; F. A. Machado et al., 7 – 9 Oct 2006. MZUSP 61056 (1, 202.2 mm SL); MZUSP 60423 (2, 255.7 – 263.5 mm SL): Nova Mutum, rio Arinos, Fervedouro, fazenda Fervedouro, 14 ° 13 ' 10 ' ’ S 56 ° 01 ' 43 ' ’ W; F. C. T. Lima et al., 18 – 19 Feb 2000. MZUSP 61133 (3, 219.3 – 247.1 mm SL); MZUSP 61132 (9, 211.9 – 434.7 mm SL); MZUSP 91506 (3, 210.0 – 247.0 mm SL): Nova Mutum, rio Arinos, 14 ° 13 ’ S, 56 ° 2 ’ W; J. M. Mendes, 1999 – 2000. MZUSP 67832 (130, 11 cs, 16.9 – 53.2 mm SL): Nova Mutum, piscicultura Buriti (stocked juveniles); H. F. Mendes, 10 Feb 2000. USNM 194317 (1, 86.5 mm SL); USNM 199204 (3, 151.3 – 194.8 mm SL); USNM 194390 (2, 170.9 – 177.5 mm SL): “ Upper Juruena-Arinos ” (no precise locality); H. Schultz, 1962. Rio Madeira basin. Rondônia: MZUSP 14028 (6, 104.9 – 130.7 mm SL): Cururu, rio Machado, c. 8 ° 13 ’ S, 62 ° 46 ’ W; M. Goulding, April 1978. UF 100614 (1, 207.6 mm SL): Rio Candeias ca. 2 km above from its mouth, c. 8 ° 38 ’ S, 63 ° 33 ’’ W; J. P. Viana, 14 May 1994. INPA 16437 (1, 112.1 mm SL): rio Jamari, diverting channel of UHE Samuel (cofferdam), 8 ° 45 ’ S, 63 ° 27 ’ W; G. M. Santos, 7 – 9 June 1988. INPA 16413 (6, 257.6 – 272.8 mm SL): rio Jamari, below Samuel dam (igapó); G. M. Santos, 27 March 1986. INPA 16457 (1, 284.8 mm SL): igarapé Japiim, c. 45 km above Samuel dam, c. 9 ° 12 ’ S, 63 ° 13 ’ W; G. M. Santos, 13 June 1985. INPA 16458 (1, 319.8 mm SL): rio Jamari, above Samuel dam; G. M. Santos, 7 Dec 1984. LBP 5146 (1, 215.0 mm SL): Ji-Paraná, rio Machado, 10 ° 46 ’ 36 ’’ S, 61 ° 55 ’ 12 ’’ W; J. Damaceno & W. Troy, 22 Aug 2007. INPA 16459 (1, 325.2 mm SL): Ariquemes, rio Jamari, below rio Canaã mouth, 9 ° 56 ’ S, 63 ° 5 ’ W; G. M. Santos, 15 Nov 1983. INPA 16363 (6, 110.5 – 148.3 mm SL): mouth of rio Guaporé, Surpresa, 11 ° 53 ’ S, 65 ° 1 ’ W; G. M. Santos, 16 Jun 1984. INPA 16356 (1, 246.0 mm SL): Pimenteiras do Oeste, rio Guaporé, 13 ° 29 ’ S, 61 ° 3 ’ W; G. M. Santos, 30 Nov 1984. CAS 68857 (1, 241.6 mm SL): Rio Guaporé, Maciel, c. 12 ° 34 ’ S, 63 ° 30 ’ W; J. D. Haseman, 26 Jul 1909. Mato Grosso: MZUSP 77402 (1, 186.8 mm SL): Panelas, rio Roosevelt, above the rapids, 9 ° 11 ’ 17 ’’ S, 60 ° 44 ’ 53 ’’ W; F. A. Machado et al., 17 – 18 July 1997. INPA 16389 (3, 161.3 – 191.1 mm SL): rio Aripuanã, Lago Genipapo; Eq. Ictiologia / INPA, 20 Aug 1976. INPA 16412 (5, 102.0 – 127.1 mm SL): rio Aripuanã, Ilha do Castanhal; Eq. Ictiologia / INPA, 20 Aug 1976. INPA 16395 (1, 283.2 mm SL): rio Aripuanã, 3 km below Cachoeira Grande, c. 10 ° 8 ’ S, 59 ° 26 ’ W; Eq. Ictiologia / INPA, 8 Nov 1976. MZUSP 103036 (1, 290.0 mm SL): Aripuanã, rio Aripuanã, below Salto de Dardanelos, 10 ° 10 ’ 6 ’’ S, 59 ° 26 ’ 50 ’’ W; F. A. Machado et al., 17 Sept 2004. MZUSP 103037 (1, 225.0 mm SL): rio Verde, trib. rio Guaporé, Brazil / Bolivia border, 14 ° 7 ’ 36 ’’ S, 60 ° 28 ’ 30 ’’ W; O. A. Cantelmo & L. Barbosa, 13 – 21 Aug 2005. MZUSP 94990 (2, 230.0 – 264.0 mm SL): Vila Bela da Santíssima Trindade, rio Guaporé, 15 ° 1 ’ S, 59 ° 49 ’ W; R. O. Mascarenhas et al., 5 Dec 2006. Rio Negro basin. Amazonas: CAS 68836 (2, 116.4 – 119 mm SL): market at Manaus; C. Ternetz, Dec 1924. MZUSP 6136 (1, 162.8 mm SL): rio Negro, above Manaus, c. 3 ° 4 ’ S, 60 ° 16 ’ W; EPA, 22 – 25 April 1967. MZUSP 59076 (3, 143.7 – 178.4 mm SL); MZUSP 59075 (1, 202.7 mm SL); MZUSP 59078 (1, 215.0 mm SL); MZUSP 59081 (1, 240.8 mm SL); MZUSP 59087 (2, 100.6 – 196.5 mm SL): Anavilhanas, Lago do Prato; G. Borges, May 1981 – June 1982. MZUSP 59084 (1, 190.0 mm SL): rio Negro, Anavilhanas, lago Xilauá (igapó); G. Borges, Aug 1981. MZUSP 56782 (3, 182.8 – 200.0 mm SL); MZUSP 59083 (4, 102.0 – 120.3 mm SL); MZUSP 59082 (1, 102.3 mm SL); MZUSP 59073 (10, 89.4 – 114.7 mm SL): rio Negro, Anavilhanas, c. 2 ° 43 ’ S, 60 ° 41 ’ W; M. Goulding, Oct 1979 – Dec 1980. MZUSP 59077 (1, 188.0 mm SL): Anavilhanas, rio Negro, Lago Camauiri (igapó): G. Borges, July 1981. MZUSP 59079 (1, 132.8 mm SL): rio Negro, Anavilhanas; G. Borges, Oct 1981. MZUSP 27333 (4, 81.1 – 96.4 mm SL): Ayrão Velho, mouth of rio Padauari, rio Negro, 2 ° 6 ’ S, 61 ° 13 ’ W; L. P. Portugal, 8 Nov 1982. MZUSP 27328 (1, 104.8 mm SL): Moura, Pedra do Gavião, rio Negro, c. 1 ° 27 ’ S, 61 ° 38 ’ W; L. P. Portugal, 13 – 14 Nov 1982. MZUSP 17658 (1, 178.0 mm SL): Rio Jauaperi (trib. rio Negro), c. 1 ° 16 ’ S, 61 ° 36 ’ W; T. R. Roberts, Nov 1968. MCP 46478 (1, 26.3 mm SL): igarapé Cambeua (trib. Rio Jauaperi), 1 ° 30 ’ 53 ’’ S, 61 ° 27 ’ 33 ’’ W; P. Petry, 28 Jan 2011. MZUSP 59074 (9, 91.6 – 113.9 mm SL): rio Negro, Cachoeira do Buiu-Açú, c. 0 ° 30 ’ S, 64 ° 50 ’ W; M. Goulding, Oct 1979. MZUSP 45526 (1, 103.3 mm SL): rio Negro, Tapera, c. 0 ° 12 ’ S, 64 ° 4 ’ W; EPA, 1 Nov 1972. MZUSP 59086 (2, 144.7 – 206.0 mm SL): rio Negro, below rio Daraá (rapids), c. 0 ° 27 ’ S, 64 ° 44 ’ W; M. Goulding, 16 Feb 1980. MZUSP 59080 (1, 285.3 mm SL): igarapé do Acará, São Pedro (igapó), c. 0 ° 21 ’ S, 66 ° 36 ’ W; M. Goulding, May 1979. LBP 6878 (2, 82.2 – 93.7 mm SL): São Gabriel da Cachoeira, rio Negro, 0 ° 8 ’ 9 ’’ S, 67 ° 05 ’ 3 ’’ W; C. Oliveira et al., 11 Aug 2008. MZUSP 91576 (3, 127.1 – 138.1 mm SL): Rio Uaupés (no precise locality); J. Chernella, no date. MZUSP 91493 (1, 226.0 mm SL): Rio Tiquié, between Caruru and Boca de Sal villages, 0 ° 16 ’ N, 69 ° 54 ’ W; M. C. Lopes, 2001 – 2002. Rio Branco basin, Roraima: MZUSP 59085 (6, 120.8 – 202.9 mm SL): Marará, rio Branco; M. Goulding, 26 Oct 1979. USNM 202716 (1, 231.4 mm SL): Rio Xeruini, c. 60 km above mouth, c. 1 ° 55 ’ S, 61 ° 55 ’ W; M. R. Brittan, 28 April 1964. MZUSP 56944 (1, 198.1 mm SL); MZUSP 59621 (3, 207.9 – 252.6 mm SL); MZUSP 56807 (10, 203.2 – 273.9 mm SL): rio Branco, cachoeira do Bem Querer, 1 ° 55 ’ N, 61 ° 1 ’ W; M. Goulding, 6 – 9 Jan 1984. MZUSP 29925 (1, 90.8 mm SL); MZUSP 56778 (1, 200.3 mm SL): igarapé 2 km above Cachoeira do Bem Querer, rio Branco; M. Goulding, 9 Jan 1984. MZUSP 29924 (2, 84.6 – 86.8 mm SL): igarapé do Bota-Panela, near Cachoeira do Bem Querer, rio Branco; M. Goulding, 9 Jan 1984. MZUSP 17756 (3, 122.9 – 134.4 mm SL): igarapé do Pau Roxo, trib. rio Uraricoera, Fazenda Canadá; T. R. Roberts, 18 Feb 1969. MZUSP 56781 (2, 211.4 – 233.9 mm SL); MZUSP 29927 (5, 107.9 – 116.7 mm SL): Igarapé do Cujobim, rio Branco, facing ilha de Maracá, c. 3 ° 22 ’ N, 61 ° 22 ’ W; M. Goulding, 13 Jan 1984. INPA 16388 (1, 169.0 mm SL): rio Mucajaí, c. 1 km below Paredão, 2 ° 56 ’ 47 ’’ N, 61 ° 34 ’ 34 ’’ W; M. Jégu & E. Ferreira, 4 Oct 1986. INPA 16394 (1, 262.6 mm SL): rio Mucajaí, c. 2 km above Paredão, 2 ° 56 ’ 20 ’’ N, 61 ° 37 ’ 14 ’’ W; M. Jégu & E. Ferreira, 3 Oct 1986. Other river systems: Brazil, Amazonas: MZUSP 99213 (1, 117.0 mm SL): Lago Urini, mouth of rio Japurá, c. 2 ° 9 ’ S, 65 ° 9 ’ W; R. B. Barthem, 30 Sept 1979. MCP 29771 (1, 121.1 mm SL): Maraã, Lago Amanã, mouth of igarapé Uxi, 2 ° 32 ’ 47 ’’ S, 64 ° 40 ’ 10 ’’ W; W. G. R. Crampton, 14 Dec 1997. MCP 29776 (1, 120.6 mm SL): Maraã, Lago Amanã, mouth of igarapé Uxi, 2 ° 37 ’ 4 ’’ S, 64 ° 40 ’ 1 ’’ W; W. G. R. Crampton, 1 Dec 1998. MCP 29774 (1, 128.1 mm SL): Maraã, Lago Amanã, mouth of rio Baré, 2 ° 28 ’ 28 ’’ S, 64 ° 43 ’ 20 ’’ W; W. G. R. Crampton, 15 Nov 1998. MCP 29775 (2, 128.9 – 131.6 mm SL): Maraã, Lago Amanã, mouth of rio Baré, 2 ° 27 ’ 23 ’’ S, 64 ° 43 ’ 35 ’’ W; W. G. R. Crampton, 13 Dec 1997. MZUSP 17592 (1, 42.5 mm SL): rio Içapó, at mouth of rio Jutaí, c. 2 ° 47 ’ S, 66 ° 49 ’ W; EPA, 22 Oct 1968. MZUSP 17530 (8, 72.2 – 134.4 mm SL): Santo Antônio do Içá, igarapé da Cachoeira, Cuiauá, trib. rio Içá, c. 3 ° 6 ’ S, 67 ° 57 ’ W; EPA, 18 Oct 1968. MNRJ 21470 (1, 146.0 mm SL): rio Queixito at mouth of rio Javari, 4 ° 25 ’ 48 ’’ S, 70 ° 15 ’ 38 ’’ W; A. Parko, 1952. Peru, Depto. Loreto: INHS 106468 (2, 97.1 – 134.4 mm SL); INHS 106470 (6, 93.5 – 119.2 mm SL): Río Nanay, Pampa Chica, 3 ° 45 ’ 1 ’’ S, 73 ° 17 ’ 0 ’’ W; M. H. Sabaj et al., 22 – 27 Jul 1997. Bolivia, Depto. Beni, Rio Madeira basin: UMMZ 204246 (2, 164.1 – 247.3 mm SL); UMMZ 204394 (1, 217.2 mm SL); UMMZ 204201 (1, 98.0 mm SL); UMMZ 204445 (2, 89.9 – 90.6 mm SL); UMMZ 204930 (1, 91.2 mm SL); UMMZ 204417 (1, 80.4 mm SL); UMMZ 204408 (2, 152.4 – 204.1 mm SL); UMMZ 204649 (1, 251.0 mm SL): Río Itenez (= Rio Guaporé), near Costa Marques, 12 ° 27 ’ S, 64 ° 14 ’ W; R. M. Bailey et al., Aug – Oct 1964. MNHN 1989 - 1429 (3, 120.2 – 146.2 mm SL): Río Itenez, Boca Machupo, c. 12 ° 27 ’ S, 64 ° 22 ’ W; L. Lauzanne & G. Loubens, no date.	en	Lima, Flávio C. T. (2017): A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae). Zootaxa 4222 (1): 1-189, DOI: 10.5281/zenodo.257769
