Brycon howesi , Lima, Flávio C. T., 2017

Lima, Flávio C. T., 2017, A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae), Zootaxa 4222 (1), pp. 1-189: 47-51

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Brycon howesi

new species

Brycon howesi  new species

( Figs. 28–29View FIGURE 28View FIGURE 29)

Chalceus opalinus  (non Cuvier): Valenciennes, in Cuvier & Valenciennes, 1850: 246 (part; “ Rio Tiquilelonha”).

Brycon insignis: Steindachner, 1877: 587  –591 (part; “ Rio Jequitinhonha”); Godinho et al., 1999: 415, 418, 421 (Rio Jequitinhonha basin; ecology).

Brycon opalinus  (non Cuvier): Géry & Mahnert, 1992: 813 –814 (part; Rio Jequitinhonha).

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).

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. 5View FIGURE 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. 6View FIGURE 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  ).

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 ( MZUSPAbout 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 12th (2) or 13th (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 22th (1), 23th (1), or 24th (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).

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.

Distribution. Endemic from the rio Jequitinhonha basin, an independent coastal river system at the state of Minas Gerais, Brazil ( Fig. 25View FIGURE 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.

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).

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 ( MNHNAbout 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, MNHNAbout MNHN A.8613 (see item “Remarks” of Brycon opalinus  , below). Although Géry & Mahnert (1992: 813) considered the specimen MNHNAbout 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.

Material examined. Holotype: MZUSPAbout MZUSP 103075View Materials (230.6 mm SL): Brazil, Minas Gerais, Araçuaí, rio Jequitinhonha, confluence with rio Araçuaí , below village of Itira , 16°45’S, 42°0’WGoogleMaps  ; J.C. Garavello, A.S. Soares, A.I. Alves & J.C. Soares, 19 Feb 1989. 

Paratypes: Brazil, Minas Gerais, rio Jequitinhonha basin: MZUSPAbout MZUSP 53803View Materials (1, 234.8 mm SL): same data as holotypeGoogleMaps  . MZUSPAbout MZUSP 53802View Materials (1, 223.1 mm SL): Leme do Prado, rio Araçuaí at Santa Rita village, 17°6’S, 42°39’W; G.B. SantosGoogleMaps  , A.S. Soares & A.I.S. Alves, 18 Oct 1987. MZUSPAbout MZUSP 55963View Materials (6, 165.8– 248.6 mm SL)  ; ZUECAbout ZUEC 6365View Materials (1, 164.9 mm SL): Coronel Murta, rio Jequitinhonha , 16°37’32’’S, 42°16’6’’W; F. Andrade, 6 Aug 2010GoogleMaps  . MZUSPAbout MZUSP 5139View Materials (6, 1 cs, 31.3–47.6 mm SL), Itaobim, rio Jequitinhonha , 16°34’S, 41°29’W; Exc. Departamento de Zoologia, 25 June 1966GoogleMaps  . ZUECAbout ZUEC 6939View Materials (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 2011GoogleMaps  . ZUECAbout ZUEC 6940View Materials (1, 143.7 mm SL): Grão Mogol, rio Itacambiruçu , 16°35’12’’S, 42°51’15’’W; T.C. Pessali & T.F. Teixeira, May 2011GoogleMaps  . ZUEC 7003 (1, 282.0 mm SL); MCP 48399 (1, 268.0 mm SL); ANSPAbout ANSP 200242View Materials (1, 232.0 mm SL): Grão Mogol, rio Itacambiruçú , 16°34’52’’S, 42°52’25’’W; T.C. Pessali & AGoogleMaps  . A. Rodrigues, Feb 2011  . ZUECAbout ZUEC 7000View Materials (1, 208.0 mm SL): Grão Mogol, rio Itacambiruçú , 16°35’50’’S, 42°50’22’’W; T.C. Pessali & AGoogleMaps  . A. Rodrigues, Feb 2011  . ZUECAbout 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 & AGoogleMaps  . A. Rodrigues, 25 Dec 2011  . ZUECAbout 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 & AGoogleMaps  . A. Rodrigues, 28 Dec 2011  . MZUSPAbout MZUSP 101506View Materials (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 2006GoogleMaps  . MZUSPAbout MZUSP 101507View Materials (3, 186.1–243.0 mm SL): same locality and collectorGoogleMaps  , 5 April 2007. MZUSPAbout MZUSP 103685View Materials (2 skel., 168.0–207.0 mm SL): same locality and collectorGoogleMaps  , July 2009. MZUSPAbout MZUSP 101508View Materials (3, 244.0–284.0 mm SL): same locality; F. Vieira & G.B. SantosGoogleMaps  , 10 Jan 2006. MZUEL 3931 (1, 130.0 mm SL): same locality; E.T. GrandoGoogleMaps  , 2 Feb 2004.

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 1989GoogleMaps  . ZUECAbout ZUEC 6969View Materials (1, 101.7 mm SL): Grão Mogol, rio Itacambiruçú , 16°35’12’’S, 42°51’15’’W; T.C. Pessali & AGoogleMaps  . A. Rodrigues, Feb 2011. ZUECAbout ZUEC 6970View Materials (2, 109.1– 138.4 mm SL): Grão Mogol, rio Itacambiruçú , 16°35’12’’S, 42°51’15’’WGoogleMaps  ; T.C. Pessali & A  . A. Rodrigues, Feb 2011. ZUECAbout ZUEC 6974View Materials (3, 101.1– 171.5 mm SL): Grão Mogol, rio Itacambiruçú , 16°34’52’’S, 42°52’25’’WGoogleMaps  ; T.C. Pessali & A  . A. Rodrigues, Feb 2011. ZUECAbout ZUEC 6999View Materials (1, 173.3 mm SL): Grão Mogol, rio Itacambiruçú , 16°34’24’’S, 42°49’46’’WGoogleMaps  ; T.C. Pessali & A  . A. Rodrigues, Feb 2011. ZUECAbout ZUEC 6367View Materials (1, 94.3 mm SL): Grão Mogol, rio Itacambiruçu , 16°35’56’’S, 42°50’8’’WGoogleMaps  ; F. Andrade, 22 Nov 2010. Imprecise localities: NMWAbout NMW 62927View Materials (1, 71.6 mm SL): “ Rio Jequitinhonha” (no specific locality); Wertheimer , no date. Paralectotype of Brycon insignis Steindachner    . MNHNAbout MNHN A.8612 (1, 295.8 mm SL): “ Rio Tiquilelonha  . A. Saint- Hilaire, 1822”. 

TABLE 7. Morphometric data of Brycon howesi (A: holotype, MZUSP 103075).

  A n Range Mean
Standard length (SL) 230.6 36 71.6–284.0 -
Percentages of standard length
Depth at dorsal-fin origin 28.6 36 25.7–33.2 29.3
Snout to dorsal-fin origin 52.9 36 46.8–56.0 52.6
Dorsal-fin base length 11.8 36 10.1–13.4 12.1
Posterior terminus of dorsal fin to adipose fin 23.8 36 21.5–25.0 23.3
Posterior terminus of dorsal fin to hypural joint 35.6 36 34.0–39.5 37.0
Snout to pelvic-fin insertion 47.5 36 47.5–53.6 49.9
Snout to anal-fin origin 64.6 36 64.4–71.7 67.7
Anal-fin base length 23.7 36 19.1–26.4 23.8
Caudal peduncle length 15.8 36 11.9–15.8 13.4
Dorsal-fin height 19.2 36 17.9–21.6 20.0
Pectoral-fin length 17.9 36 16.1–20.9 19.3
Pelvic-fin length 14.1 36 14.1–16.8 15.2
Caudal peduncle depth 8.5 36 8.1–9.3 8.7
Head length 26.9 36 25.3–31.7 28.6
Percentages of head length
Head height 70.9 36 65.6–76.9 71.3
Snout length 30.8 36 26.8–32.3 29.9
Upper jaw length 44.0 36 42.7–50.2 45.9
Horizontal eye diameter 21.4 36 19.0–27.3 22.7
Post-orbital length 48.6 36 45.8–53.2 50.3
Least interorbital width 34.5 36 28.6–37.1 33.0

Museu de Zoologia da Universidade de Sao Paulo


Museum National d'Histoire Naturelle


Museu de Zoologia da Universidade Estadual de Campinas


Academy of Natural Sciences of Philadelphia


Naturhistorisches Museum, Wien














Brycon howesi

Lima, Flávio C. T. 2017



Pugedo 2016: 345


Brycon devillei

Azevedo 2011: 807


Brycon insignis:

Godinho 1999: 415Steindachner 1877: 587


Brycon opalinus

Gery 1992: 813


Chalceus opalinus

Cuvier 1850: 246