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Hyphessobrycon langeanii was originally described as endemic to the upper Araguaia river basin, Brazil. However, our analysis of several Hyphessobrycon specimens collected in the Correntes river basin and another tributary of the Itiquira River in Mato Grosso state (both belonging to the Paraguay river basin) reveals the first verified record of H. langeanii from this basin and from the states of Mato Grosso do Sul and Mato Grosso. The objective of this study is to provide a novel record of H. langeanii from the basin of the Paraguay River and from the state of Mato Grosso do Sul. We compare the newly discovered populations with the populations in the Araguaia river basin, and we also provide a brief discussion on the biogeography of this species.

Hyphessobrycon bussingi, new species (Figs. 1–6, Tab. 1) urn:lsid:zoobank.org:act: DDDE4FB2-D3E6-4019-88D3-FF734DE6AFD3 Hyphessobrycon panamensis (not Durbin)—. Behre, 1928: 318 [list of species, Western Panama; distribution]—. Bussing, 1966:218 [in part; lots ANSP 99920 and ANSP 99932]—. Bussing, 1987: 85 [Costa Rica; brief description, photo, distribution]—. Bussing, 1998: 105-108 [Costa Rica; diagnosis in key, brief description, photo, distribution]—. Lima et al., 2003: 139 [in part; listed; Costa Rica]—. Angulo et al., 2013: 992 [list of species, Costa Rica; distribution]. Holotype. UCR 3302-001, 33.7 mm SL, male (rd): Costa Rica, Limón, río Cocolis, 3.5 km southeast of Shiroles, on road Bratsi-Shiroles, 09°34’05’’N, 82°56’4.92’’W; W. A. Bussing ichthyology team, 06 Oct 1979. Paratypes. All from Costa Rica, Limón. Río Sixaola basin: MZUSP 125290, 15, 27.5 –32.0 mm SL, San Miguel, Crique Azul on finca Los Laureles de ANAI, 09°33’50’’N 82°40’39.4’’W; W. McLarney, 31 Mar 1987 NUP 22293, 9, 27.2–34.0 mm SL, stream tributary of río Gandoca, finca Elida Morales, 09°34’10’’N 82°37’19.7’’W; W. McLarney, 18 Mar 1983; UCR 3240001, 39, 24.4–34.0 mm SL, same data as holotype; UCR 3241001, 20, 26.4–36.0 mm SL, Mojoncito, Bratsi, stream Shiragri, 09°30’34’’N 83°00’01’’W; A. E. Corrales & C. Jiménez, 05 Jul 1982; UCR 3242001, 30, 26.2–29.3 mm SL, stream Ñiabri, 09°29’25’’N 82°58’24.6’’W; A. Corrales, C. Jiménez & A. Román, 12 Jun 1982. Non-type material. Río Sixaola basin: FMNH 144016, 8 (2), laguna Mata de Limón, 9°34’5.06”N 82°37’46.03”W; C. McMahan, W. Ludt, A. Angulo & M. Mafla, 09 Nov 2019; UCR 129917, 119, same data as holotype; UCR 1395007, 74, stream Ñiabri, 09°29’25’’N 82°58’24.6’’W; A. Corrales, C. Jiménez & A. Román, 12 Jun 1982; UCR 1408001, 52, Mojoncito, Bratsi, stream Shiragri, 09°30’34’’N 83°00’01’’W; A. E. Corrales & C. Jiménez, 05 Jul 1982; UCR 1838005, 20, stream tributary of río Gandoca, finca Elida Morales, 09°34’10’’N 82°37’19.7’’W; W. McLarney, 18 Mar 1983; UCR 1881002, 28, San Miguel, Crique Azul on finca Los Laureles de ANAI, 09°33’50’’N 82°40’39.4’’W; W. McLarney, 31 Mar 1987. Panama, Bocas del Toro: ANSP 99932, 2, 35.6–36.1 mm SL, backwaters and overflowed banks of río Guarumo at Chiriquicito, 8°57’N 82°11’W; H.G. Loftin & R. W. Yerge, 19 Apr 1962; ANSP 104149, 31 (5, 32.7–35.1 mm SL), río Changuinola upstream about 10 mi to first big rapid, ʽCorriente Grandeʼ, at creek & backwater, 9°30’N 82°43’W; H. G. Loftin, 30 Sep 1962. Costa Diagnosis. Hyphessobrycon bussingi differs from all congeners, except from H. compressus, H. diancistrus Weitzman & Palmer, H. otrynus Benine & Lopes, and those belonging to the H. panamensis species-group, by the presence of one large bony hook per fin ray, anterodorsally oriented and surrounded by a fleshy area, on last unbranched and first branched anal-fin rays of mature males (vs. absence of hooks, or presence of small and numerous hooks per fin ray). Hyphessobrycon bussingi can be distinguished from H. compressus by the absence of a black blotch on the dorsal fin (vs. presence), by the presence of predorsal series (vs. absence), and by presenting 32–34 scales in longitudinal series (vs. 41–48). It can be distinguished from H. diancistrus and H. otrynus by presenting caudal fin hyaline (vs. dark-brown blotches on caudal-fin lobes in H. diancistrus; distal third of caudal fin darkened in H. otrynus). It differs from H. columbianus by the absence of a silvery-blue iridescent coloration on dorsal region of flanks in life (vs. presence), and by a lower body depth (33.7–42.6% SL vs. 46.0–50.3% SL). It can be distinguished from H. savagei by presenting the second humeral blotch less conspicuous than the first, extending one or two longitudinal scale rows below lateral line, with posterior margin as a concentration of melanophores concentrated along longitudinal midlateral stripe (vs. humeral blotches evenly pigmented, extending two to four longitudinal scale rows below lateral line, with contours and margins always well-defined). It can be distinguished from H. panamensis by a higher number of branched anal-fin rays (23–27, rarely 22, mode = 24 vs. 19–23, modes = 21 and 22), and by presenting 6–7 (mode = 6½) scale rows between lateral line and dorsal-fin insertion (vs. 7–8, mode = 7). It can be distinguished from Hyphessobrycon condotensis by presenting 8–9, rarely 7 (mode = 8) gill rakers on epibranchial 1 (vs. 5–7, mode = 7), and higher dorsal- (27.8–41.2% SL vs. 27.7–29.7% SL) and pelvic-fin length (20.4–24.5% SL vs. 17.3–20.5% SL) in mature males. It differs from Hyphessobrycon daguae by presenting a well-defined first humeral blotch (vs. a single diffuse humeral blotch). Description. Morphometric data in Table 1. Body compressed, greatest body depth slightly anterior vertical through dorsal-fin origin. Dorsal profile of body convex from upper lip to vertical through anterior nostril; straight from this point to distal tip of supraoccipital process. Slightly convex from tip of supraoccipital process to dorsal-fin origin; straight from this point to adipose-fin origin; and straight or slightly concave along caudal peduncle. Ventral profile of body convex from lower lip to anal-fin origin; straight or slightly convex along anal-fin base; and slightly concave along caudal peduncle. Jaws equal, mouth terminal. Premaxillary teeth in two rows; outer row with 1(1), 2*(18), 3(21), or 4(13) tricuspid teeth; inner row with 4(1), 5*(44), or 6(6) tri- to pentacuspid teeth. Posterior margin of maxilla at vertical through anterior margin of pupil. Maxilla with 1(4), 2*(28), or 3(22) conical, tri- to pentacuspid teeth. Dentary with 4(21), 5*(26), or 6(1) large tri- to pentacuspid teeth, gradually decreasing in size, followed by five conical and abruptly smaller teeth. Central cusp longer than lateral ones. Scales cycloid. Lateral line incompletely pored with 6(4), 7(9), 8(27), 9*(21), 10(2), or 11(1) perforate scales. Longitudinal series, including perforate scales, with 32(2), 33(31), or 34*(15). Scale rows between dorsal-fin origin and lateral line 6(16), 6½*(9), or 7(37). Scale rows between lateral line and pelvic-fin insertion 4(2), 4½ (9), 5*(42), or 5½(8). Predorsal series irregular*, uni or biserial, with 10(22), 11(15), or 12(4) on middline. Single row of scales covering base of anteriormost anal-fin rays. Circumpeduncular scales 13*(15) or 14(20). Dorsal-fin rays ii,8,i(7) or ii,9*(46). First dorsal-fin pterygiophore located between neural spines of ninth and 10 th (1) vertebrae. Adipose fin present. Pectoral-fin rays i,11*(23) or 12(31), distal tip of rays reaching pelvic-fin origin. Pelvic-fin rays i,6(3) or 7*(52), distal tip surpassing anal-fin origin. Anal-fin rays iii(2) or iv*(51), 22(1), 23(6), 24*(23), 25(25), 26(11), or 27(1). Anal fin of mature males with large bony hooks. Caudal fin bifurcate, lobes approximately of same size, i,9/8,i rays. First gill arch with 7(2), 8(44), or 9*(10) rakers on epibranchial, 1*(55) on intermediate cartilage, 11(1), 12(23), or 13*(32) on cerato+hipobranchial. Branchiostegal rays 4. Total vertebrae (32), supraneurals 4(1) (Fig. 2). Color in alcohol. Background color light brown. Dorsal region of head and middorsal region of body with dense concentration of melanophores. Region above pupil darkened. Infraorbitals and opercle beige, with few scattered melanophores, evenly distributed. Opercle retaining some guanine pigmentation. Lower portion of maxilla and gular region with few scattered melanophores, slightly concentrated on upper half. Two dark-brown humeral blotches, vertically elongated, separated by light-beige area. First humeral blotch conspicuous, extending three to four longitudinal scale rows above and one or two below lateral line. Second humeral blotch usually inconspicuous, with melanophores equally distributed, four to five longitudinal scale rows above and one or two below lateral line; posterior margin with concentration of melanophores concentrated along longitudinal midlateral stripe. Flanks with melanophores evenly distributed, except on antero-ventral half, with few or without melanophores. Posterior margin of scales on superior third of flanks with concentration of melanophores. Brownish to silvery longitudinal midlateral stripe, from second humeral blotch to caudal peduncle, more conspicuous on posterior region. Fins hyaline, with slight concentration of melanophores on distal half of dorsal fin, distal third of anal fin, and median caudal-fin rays. Abdominal region, from pectoral-fin origin to pelvic-fin end, lacking melanophores. Color in life. Background color pale beige to silvery. Dark markings as in preserved specimens. Abdominal and region above anal fin reddish. Pelvic fin and distal margin of anal fin reddish. Adipose- and dorsal fins reddish or yellowish during the reproductive period (Bussing, 1987, 1998, and Fig. 3). Distribution. Hyphessobrycon bussingi is known from the río Sixaola basin, Atlantic coast of Costa Rica, and río Changuinola, río Guarumo, and río San San, Panama (Fig. 4, and 21). Sexual dimorphism. Mature males with one large bony hook per fin ray, anterodorsally oriented and surround- ed by a fleshy area, from the last unbranched (on fourth or fifth segments) to the fifth branched (varying on fifth to seventh segments) anal-fin rays of mature males (Figs. 5a and 6a). One specimen presented two large bony hooks on last unbranched and first branched anal-fin rays. Very small hooks, difficult to discern, can be present on distal segments of most anterior anal-fin rays. Dorsal- and pelvic-fin length of males slightly longer than in females. Males with distal margin of anal fin almost straight, and females with distal margin of anal fin falcate, with anteriormost anal-fin rays longer than posterior ones. Ecology and habitat. Hyphessobrycon bussingi feeds on terrestrial insects, and lives in rivers and streams with moderate flow, in altitudes ranging from 40 to 60 meters a.s.l. and with water temperatures ranging from 24–26ºC (Bussing, 1998, as H. panamensis). Etymology. The specific epithet honors William A. Bussing (1933-2014), in recognition of all his contributions to the knowledge of the ichthyofauna of Costa Rica. A genitive noun. Remarks. Hyphessobrycon bussingi was previously incorrectly identified as H. panamensis in Costa Rica and in the province of Bocas del Toro, Panama (at the border with Costa Rica) (e.g. Bussing, 1987, 1998; Angulo et al., 2013). However, both species can be diagnosed by the characters mentioned in the Diagnosis section, above. Hyphessobrycon columbianus Zarske & Géry, 2002 (Figs. 7–8, Tab. 2) Hyphessobrycon columbianus Zarske & Géry, 2002: 22 [original description, type locality: “ Kolumbien, Darien, kleiner Bach etwa 6 km flussaufwärts von Acandi, Einzugsgebiet des Rio Acandi, Atlantikküste”]—.Lima et al., 2003: 135 [listed; distribution]—. Zarske, 2003: 17 [type catalogue]—. Maldonado-Ocampo et al. 2008: 168 [Colombia; Pacific slope rivers; listed]—. García-Alzate et al., 2010: 55 [diagnosis of H. sebastiani; material examined: río Guati, Acandi, Chocó]—. Maldonado-Ocampo et al., 2012: 117 [Colombia, Chocó; photograph; distribution] —. García-Alzate et al., 2013: 182 [diagnosis of H. chocoensis; material examined: río Guati, Acandi, Chocó; key to species]—. DoNascimiento et al., 2017: 49 [occurrence in Colombia]. Diagnosis. Hyphessobrycon columbianus differs from all congeners by the presence of a silvery-blue iridescent coloration on dorsal region of flank in life (vs. absence of a similar color pattern). It can be further distinguished from all congeners, except from H. compressus, H. diancistrus, H. otrynus, and those belonging to the H. panamensis species-group, by the presence of one large bony hook per fin ray, antero-dorsally oriented and surrounded by a fleshy area, on last unbranched and first branched anal-fin rays of mature males (vs. absence of hooks, or presence of small and numerous hooks per fin ray). It also differs from H. compressus by the absence of a black blotch on the dorsal fin (vs. presence), by the presence of predorsal series (vs. absence), and by presenting 32–34 scales in longitudinal series (vs. 41–48). It can be distinguished from H. diancistrus and H. otrynus by presenting the caudal fin hyaline (vs. dark blotches on caudal-fin lobes in H. diancistrus; distal third of caudal fin darkened in H. otrynus). Additionally, it can be distinguished from the remaining species of the H. panamensis species-group by a higher body depth (46.0–50.3% SL vs. 31.8–46.5% SL). Description. Morphometric data in Table 2. Body compressed, greatest body depth slightly anterior to vertical through dorsal-fin origin. Dorsal profile of body convex from upper lip to vertical through anterior nostril; slightly convex from this point to basis of supraoccipital; slightly concave to almost straight from base- to distal tip of supraoccipital process. Convex from tip of supraoccipital process to dorsal-fin origin; straight or slightly convex from this point to adipose-fin origin; and straight or slightly concave along caudal peduncle. Ventral profile of body convex from lower lip to pelvic-fin origin; straight or slightly convex from this point to anal-fin origin; straight to slightly convex along anal-fin base; and straight or slightly concave along caudal peduncle. Jaws equal, mouth terminal. Premaxillary teeth in two rows; outer row with 1(1), 2(2), 3(5), 4*(2) tricuspid teeth; inner row with 5*(11) tri- to pentacuspid teeth. Posterior margin of maxilla slightly posterior at vertical through anterior margin of pupil. Maxilla with 2(4) or 3*(7) conical and tricuspid teeth. Dentary with 4(3) or 5*(6) large penta and tricuspid teeth, gradually decreasing in size, followed by 10 conical and abruptly smaller teeth. Central cusp more developed than lateral ones. Scales cycloid, radii 2–9. Lateral line incompletely pored with 7*(1), 8(1), 9(1), 10(3), 11(2), 13(2), 16(1), or disrupted with total of 20(1) perforate scales. Longitudinal series, including perforate scales, with 32*(6), 33(5), or 34(1). Scale rows between dorsal-fin origin and lateral line 6(4), 7*(4), or 7½(4).). Scale rows between lateral line and pelvic-fin insertion 5*(5), 5½(1), 6(3), or 6½(4). Predorsal series with 9*(3), 10(8), or 11(2) scales. Single row of six scales covering base of anteriormost anal-fin rays. Circumpeduncular scales 14*(11). Dorsal-fin rays ii,9(12) or 10*(1). Adipose fin present. Pectoral-fin rays i,10*(5), 11(7), or 12(1), distal tip of rays reaching pelvic-fin origin. Pelvic-fin rays i,6(2) or 7*(11), distal tip reaching anal-fin origin. Anal-fin rays iii* or iv(2), 22(3), 23(7), or 24*(1). Anal fin of mature males with large bony hooks. Caudal fin bifurcate, lobes approximately of same size, i,9/8,i*(3) rays. First gill arch with 5(2), 6(7), or 7(2) rakers on epibranchial, 1(11) on intermediate cartilage, 8(1), 9(4), or 10(6) on ceratobranchial + hipobranchial. Branchiostegal rays 4. Color in alcohol. Background coloration light brown. Dorsal region of head and middorsal region of body with dense concentration of melanophores. Region above pupil darkened. Infraorbitals and opercle beige, with scattered melanophores more concentrated on upper half. Opercle with some guanine pigmentation. Distalmost portion of maxilla and gular region with few scattered melanophores, slightly more concentrated on upper half. Two darkbrown humeral blotches, vertically elongated, separated by light-beige area. First humeral blotch conspicuous, with melanophores more concentrated on central region, extending four longitudinal scale rows above and one or two below lateral line. Second humeral blotch usually inconspicuous, with melanophores evenly distributed, extending four to five longitudinal scale rows above and one below lateral line; posterior margin with concentration of melano- phores concentrated along longitudinal midlateral stripe. Body sides with melanophores evenly distributed, except on anteroventral half, with few or without melanophores. Posterior margin of scales on superior half of flank with concentration of melanophores. Brownish to silvery longitudinal midlateral stripe, from second humeral blotch to caudal peduncle, more conspicuous on posterior region. Fins hyaline; distal half of dorsal fin, and base distal third of anal fin with more numerous melanophores. Abdominal region, from pectoral-fin origin to pelvic-fin end, with few melanophores. Color in life. (based in Zarske & Géry, 2002, fig. 2, p. 25). Background color greenish golden. Dark markings as in preserved specimens, except for humeral blotches inconspicuous. Dorsal region of body greenish-golden. Upper half of head brown; lower half of head golden. Iris golden. Dorsal region of flank with silvery-blue iridescent coloration. Region above anal fin reddish. Abdominal region whitish-yellow. Dorsal fin reddish-yellow. Pectoral and pelvic fins hyaline. Anal fin reddish-orange. Caudal fin reddish, with distal margin darkened. Distribution. Hyphessobrycon columbianus is known from the río Acandi (Caribbean Sea), in Colombia, and drainages of the province of San Blas, southeast Panama (Fig. 8, and 21). Sexual dimorphism. Mature males with one large bony hook per fin ray, antero-dorsally oriented and surrounded by fleshy area, from the last unbranched to the third branched anal-fin ray of mature males. Very small hooks, which are difficult to discern, can be present on distal segments of most anterior anal-fin rays. In addition, males present a deeper body than females (Zarske & Géry, 2002). Material examined. 24 specimens (25.2–45.7 mm SL). Type material. MTD-F 25497, holotype, 45.7 mm SL: Colombia, Chocó, Darien region, small stream about 6 kilometers downstream from Acandi, entry of the río Acandi, Atlantic versant; Bork & Machnik leg., 20 Mar 1995. MTD-F 25498, paratype, 40.9 mm S

Hyphessobrycon condotensis Regan, 1913 (Figs. 9���10, Tab. 3) Hyphessobrycon condotensis Regan, 1913: 465 [original description, type locality: ���Rio Condoto (���) and the Rio San Juan���, Colombia]���. Eigenmann, 1922: 141 [as a synonym of H. panamensis]���. Zarske & G��ry, 2002: 26 [revalidation of H. condotensis; key to species]���.Lima et al., 2003: 135 [listed; distribution]���. Maldonado-Ocampo et al. 2008: 168 [listed; Colombia; Pacific slope rivers; distribution]���. Garc��a-Alzate et al., 2010: 55 [comparison with H. sebastiani; material examined]���. Maldonado-Ocampo et al., 2012: 118 [Colombia, Choc��; photograph; short description; distribution] ���. Garc��a-Alzate et al., 2013: 182 [recorded at Colombia; diagnosis of H. sebastiani; material examined; key to species]���. Garc��a-Alzate et al., 2015: 222 [material examined: r��o Baud��, r��o Condoto and r��o San Juan]���. DoNascimiento et al., 2017: 49 [occurrence in Colombia]. Hyphessobrycon panamensis (not Durbin)���. Eigenmann, 1918: 186 [in part; specimens from r��os Atrato, San Juan, and Magdalena basins, Colombia]���. Eigenmann, 1920a: 17 [in part; specimens from r��o Atrato, r��o San Juan, and r��o Magdalena basins]���. Eigenmann, 1920b: 15 [in part; specimens from r��o Atrato]���. Eigenmann, 1920c: 11 [Western Colombia, r��o Atrato and r��o San Juan basins; distribution]���. Eigenmann, 1920d: 29 [Colombia, lower r��o Magdalena basin; distribution]���. Eigenmann, 1922: 141 [in part; specimens from Colombia: Calamar, Calima, Certegui, Condoto, Istmina, Managru, Truando]���. Dahl, 1971: 140 [in part; specimens from r��os Atrato, San Juan and lower Magdalena, Colombia]���. Weitzman & Palmer, 1997: 228 [in part; specimens from r��os Atrato, San Juan and lower Magdalena, Colombia]���.Lima et al., 2003: 139 [in part; r��os Atrato, San Juan and lower Magdalena, Colombia]. Hyphessobrycon sebastiani Garc��a-Alzate et al., 2010: 55 [original description, type locality: ��� Colombia, Pacific versant, Choc��, Istmina, San Juan River drainage, Patecucho Creek, approximately 5��9���N & 76��40���W ���]���. Maldonado-Ocampo et al., 2012: 119 [Colombia, Choc��; photograph; short description; distribution; biological data]���. Garc��a-Alzate et al., 2013: 182 [diagnosis in key; material examined: r��o Condoto and r��o San Juan]���. DoNascimiento et al., 2017: 49 [list of species, Colombia]. NEW SYNONYM. Diagnosis. Hyphessobrycon condotensis differs from all congeners, except from H. compressus, H. diancistrus, H. otrynus, and those belonging to the H. panamensis species-group, by the presence of one large bony hook per fin ray, antero-dorsally oriented and surrounded by a fleshy area, on last unbranched and first branched anal-fin rays of mature males (vs. absence of hooks, or presence of small and numerous hooks per fin ray). Hyphessobrycon condotensis can be distinguished from H. compressus by the absence of a black blotch on dorsal fin (vs. presence), by the presence of predorsal series (vs. absence), and by presenting 31���35 scales in longitudinal series (vs. 41���48). It can be distinguished from H. diancistrus and H. otrynus by presenting caudal fin hyaline (vs. dark blotches on caudal-fin lobes in H. diancistrus; distal third of caudal fin darkened in H. otrynus). It differs from H. columbianus by the absence of a silvery-blue iridescent coloration on dorsal region of flank in life (vs. presence), and by the lower body depth (34.7���46.5% SL vs. 46.0���50.3% SL). It can be distinguished from H. savagei by presenting the second humeral blotch less conspicuous than the first, extending one or two longitudinal scale rows below lateral line, with posterior margin as a concentration of melanophores concentrated along longitudinal midlateral stripe (vs. humeral blotches equally pigmented, extending two to four longitudinal scale rows below lateral line, with contours and margins always well-defined). It can be distinguished from H. panamensis by the higher number of branched anal-fin rays (23���27, rarely 22, mode = 24 vs. 19���23, modes = 21 and 22), by presenting 5���7 (mode = 6) scale rows between lateral line and dorsal-fin insertion (vs. 7���8, mode = 7), and 4�����6 (mode = 5) scale rows between lateral line and pelvic-fin insertion (vs. 6, rarely 5��, mode = 6). It can be distinguished from Hyphessobrycon bussingi by presenting 5���7 (mode = 7) gill rakers on epibranchial 1 (vs. 8���9, rarely 7, mode = 8), and lower dorsal- (27.7���29.7% SL vs. 27.8���41.2% SL) and pelvic-fin length (17.3���20.5% SL vs. 20.4���24.5% SL) in mature males. It differs from Hyphessobrycon daguae by presenting a well-defined first humeral blotch (vs. a single diffuse humeral blotch). Description. Morphometric data in Table 3. Body compressed, greatest body depth at vertical through dorsalfin origin. Dorsal profile of body convex from upper lip to vertical through anterior nostril; straight or slightly concave from this point to distal tip of supraoccipital process, and slightly convex from latter point to dorsal-fin origin; straight or slightly convex from this point to adipose-fin origin; and straight along caudal peduncle. Ventral profile of body convex from lower lip to pelvic-fin origin; straight or slightly convex from this point to anal-fin origin; straight along anal-fin base; and straight or slightly concave along caudal peduncle. Jaws equal, mouth terminal. Premaxillary teeth in two rows; outer row with 2(4), 3(17), 4*(22), or 5(2) tricuspid teeth; inner row with 4(2), 5*(41), or 6(2) tri- to pentacuspid teeth. Posterior margin of maxilla at vertical through anterior margin of pupil. Maxilla with 1(5), 2(29), or 3*(10) tricuspid teeth. Dentary with 3(1), 4*(25), or 5(15) large, tri- to pentacuspid teeth, gradually decreasing in size, followed by conical and abruptly smaller teeth. Central cusp longer than lateral ones. Scales cycloid. Lateral line incompletely pored, with 6(3), 7(1), 8(5), 9(7), 10(8), 11(6), 12(6), 13(2), or 17*(1) or disrupted with total of 22(1) perforate scales. Longitudinal series, including perforate scales, with 31(2), 32(4), 33*(16), 34(10), or 35(3). Scale rows between dorsal-fin origin and lateral line 5(3), 6*(28), 6��*(1), or 7(10). Scale rows between lateral line and pelvic-fin insertion 4��(3), 5(30), 5��*(8), or 6(5). Predorsal series with 10(22), 11(14), or 12(2) scales. Single row of seven scales covering base of anteriormost anal-fin rays. Circumpeduncular scales 13(5) or 14(32). Dorsal-fin rays ii,8,i(7) or ii,9*(32). Adipose fin present. Pectoral-fin rays i,10(3), 11*(26), or 12(7), distal tip of rays reaching pelvic-fin origin. Pelvic-fin rays i,6(1) or 7*(36), distal tip reaching anal-fin origin. Anal-fin rays iii*(4) or iv(36), 22(3), 23(10), 24(19), 25(14), 26*(4), or 27(1). Anal fin of mature males with large bony hooks. Caudal fin bifurcate, lobes approximately of same size, with i,9/8,i(1) rays. First gill arch with 5(1), 6(9), or 7(14) rakers on epibranchial, 1(24) on intermediate cartilage, 10(3), 11(19), or 12(4) on cerato+hipobranchial. Branchiostegal rays 4. Color in alcohol. Background coloration light brown. Dorsal region of head and middorsal region of body with dense concentration of melanophores. Region above pupil darkened. Infraorbitals and opercle beige, with few scattered melanophores, more concentrated on upper half. Guanine pigmentation present on opercle. Lower portion of maxilla and gular region with few scattered melanophores. Two dark-brown humeral blotches, vertically elongated, separated by light-beige area. First humeral blotch conspicuous, with melanophores more concentrated on central area, extending three to four longitudinal scale rows above and two to three below lateral line. Second humeral blotch usually inconspicuous, with melanophores evenly distributed, extending three to four longitudinal scale rows above and one or two below lateral line; posterior margin with concentration of melanophores concentrated along longitudinal midlateral stripe. Body sides with melanophores evenly distributed, except on antero-ventral half, with few or without melanophores. Posterior margin of scales on superior half of flanks with concentration of melanophores. Brownish to silvery longitudinal midlateral stripe, from second humeral blotch to caudal peduncle, more conspicuous on posterior region. Fins hyaline, with slight concentration of melanophores on proximal half of dorsal fin, distal third of anal fin. Abdominal region, from pectoral-fin origin to pelvic-fin end, with few scattered melanophores. Distribution. Hyphessobrycon condotensis is known from r��o Atrato, r��o Baud��, lower r��o Magdalena and r��o San Juan basins, in Colombia (Fig. 8). Sexual dimorphism. Mature males with one large bony hook per fin ray, anterodorsally oriented and surrounded by fleshy area, from the last unbranched (on the fifth segment) up to fourth branched (on the sixth segment) anal-fin ray of mature males. Very small hooks, difficult to discern, can be present on distal segments of most anterior anal-fin rays. Dorsal-fin length of males slightly longer than in females. Ecology and habitat. Hyphessobrycon condotensis feeds mainly on insects, but some algae were also found in stomach contents (Garc��a-Alzate et al., 2010, as H. sebastiani). Remaks. Hyphessobrycon condotensis was described from the r��o Condoto, a tributary to the r��o San Juan. In the original description, Regan (1913) cited that this species was collected in the r��o Condoto by H.G.F. Spurell, and in the r��o San Juan by G. Palmer. Recently, Garc��a-Alzate et al. (2010) described H. sebastiani, also from the r��o San Juan basin, but these authors did not include H. condotensis and H. panamensis in the diagnosis of the species, comparing them only in a Principal Component Analysis (PCA), in which the measurements that most influenced the ordination were body depth, anal- and pelvic-fin length. However, the examination of the holotype and topotypes of H. sebastiani, combined with the original description and the examination of the type series of H. condotensis undertook during the present study demonstrated that all these data overlap (Table 3). Garcia-Alzate et al. (2013) provided a key to the species of Hyphessobrycon from the Pacific coastal drainages of Colombia and mentioned among the diagnostic characters of H. condotensis the presence of only a single humeral blotch, and 11 perforate scales on the lateral line. However, the syntypes of H. condotensis present two humeral blotches (even if inconspicuous), and 10���17 perforate scales on lateral line, characters also found in the type-series of H. sebastiani. Therefore, H. sebastiani is herein proposed as a junior synonym of H. condotensis. The type series of Hyphessobrycon condotensis is composed of three syntypes (BMNH 1913.10.1.19-21). The lectotype herein designated is the larger specimen (33.0 mm SL, Fig. 9), and the remaining syntypes become thus paralectoytpes, as ruled by the article 74 of ICZN (1999). Material examined. 57 specimens (14.4���40.0 mm SL). Type material. BMNH 1913.10.1.19, lectotype of Hyphessobrycon condotensis, 33.0 mm SL: Colombia, Choc��, r��o Condoto and r��o San Juan; H.G.F. Spurell & G. Palmer; BMNH 1913.10.1.20-21, paralectotypes of Hyphessobrycon condotensis, 2, 14.4 ��� 30.0 mm SL, same data as lectotype; IUQ 1942, holotype of Hyphessobrycon sebastiani, 38.4 mm SL: Colombia, Choc��, Istmina, creek Patecucho, tributary of r��o San Juan, 05��09���N 76��40���W; T. Silirio, 07 Aug 2002. Non-type material. Colombia, Choc��, r��o Atrato basin: IAvH 6497, 3, 25.1 ��� 29.1 mm SL; IAvH 6499, 1, 31.9 mm SL; IAvH 6500, 6, (1, 27.7 mm SL); IAvH 6504, 6, (2, 23.0 ��� 28.8 mm SL); IAvH 6514, 6, (3, 28.3 ��� 32.0 mm SL); IAvH 6515, 3, 29.1 ��� 33.4 mm SL; IAvH 6516, 1, 27.2 mm SL; IAvH 6517, 3, (2, 18.4 ��� 30.0 mm SL), Yuto, stream Do��a Josefa, 05��31���N 76��37���W; Chaverra & Cuesta, 21 Jun 2004; IAvH 6498, 5, 26.2 ��� 33.4 mm SL; IAvH 6501, 5, (2, 28.5 ��� 32.2 mm SL); IAvH 6502, 1, 35.8 mm SL; IAvH 6513, 1, 33.8 mm SL; IAvH 6526, 1, 36.0 mm SL; IAvH 11241, 3, 36.2 ��� 40.0 mm SL; IAvH 11246, 2, 26.3 ��� 28.6 mm SL; IAvH 11247, 4, 27.0 ��� 32.7 mm SL, r��o Yuto, 05��31���N 76��39���W; Toldillo & Cuesta, 26 Jun 2004; IAvH 7218, 1, 25.9 mm SL, Ungu��a, r��o Ungu��a, below the mouth of the aqueduct, 08��03���35,6������N 77��07���20,8������W; J. Maldonado-Ocampo et al., 24 Jul 2005. ICNMNH 1574, 10 (2, 20.8 ��� 22.3 mm SL), Quibdo, vereda Sanceno, r��o Atrato, tributary of r��o Cienaga, 05��41���N 76��39���W; C. Rom��n-Valencia, 22 Jan 1988; ICNMHN 131, 15 (3, 20.1 ��� 27.8 mm SL), stream in front of Amparraid��, r��o Baud��, 05��42���N 77��01���W; G. Dahl, Nov 1959; ICNMHN 2220, 5 (2, 29.2 ��� 31.1 mm SL), stream on road Quibdo-Itsmna, 05��34���N 76��39���W; G. Galv��s & M. Camargo, Mar 1994; UTCH 0104, 5 (1, 31.1 mm SL) stream Chaparraido, 05��37���N 76��38���W; J. Yair-Casas, O. Carrasca, 6 Feb 1994. Choc��, r��o San Juan basin: ICNMHN 205, 49 (4, 13.9 ��� 33.4 mm SL), stream Camperro, tributary of r��o San Juan, 05��04���N 76��43���W; G. Dahl, 05 Nov 1959; ICNMHN 2262, 9 (6, 28.3 ��� 34.7 mm SL), stream Profundo, tributary of r��o San Juan, 05��11���N 76��36���W; G. Galvis & M. Camargo, Mar 1994., Published as part of Ota, Renata R., Carvalho, Fernando R. & Pavanelli, Carla S., 2020, Taxonomic review of the Hyphessobrycon panamensis species-group (Characiformes: Characidae), pp. 401-436 in Zootaxa 4751 (3) on pages 412-415, DOI: 10.11646/zootaxa.4751.3.1, http://zenodo.org/record/3714610, {"references":["Regan, C. T. (1913) The fishes of the San Juan River, Colombia. Annals and Magazine of Natural History, 12 (71), 462 - 473.","Eigenmann, C. H. (1922) The fishes of Western South America [Part I]: The fresh-water fishes of Northwestern South America, includind Colombia, Panama, and the Pacific slopes of Ecuador and Peru, together with and appendix upon the fishes of the rio Meta in Colombia. Memoirs of the Carnegie Museum, 9 (1), 1 - 346.","Zarske, A. & Gery, J. (2002) Der Blaurote Kolumbien-Salmler. Hyphessobrycon columbianus n. sp. - ein neuer Salmler (Teleostei; Characiformes, Characidae) aus dem kolumbianischen Darien. Das Aquarium, 36 (391), 22 - 30.","Maldonado-Ocampo, J. A., Vari, R. P. & Usma, J. S. (2008) Checklist of the freshwater fishes of Colombia. Biota Colombiana, 9 (2), 143 - 237.","Garcia-Alzate, C. A., Roman-Valencia, C. & Taphorn, D. C. (2010) A new species of Hyphessobrycon (Teleostei: Characiformes: Characidae) from the San Juan River drainage, Pacific versant of Colombia. Zootaxa, 2349, 55 - 65. http: // doi. org / 10.11646 / zootaxa. 2349.1.4","Maldonado-Ocampo, J. A., Usma, J. S., Villa-Navarro, F. A., Ortega-Lara, A., Prada-Pedreros, S., Jimenez, L. F., Karamillo-Villa, U., Arango, A., Rivas, T. & Sanchez, G. C. (2012) Peces dulceacuicolas del Choco biogeografico de Colombia. WWF Colombia, IAvH, AU-NAP, Pontificia Universidad Javeriana, Bogota (DC).","Garcia-Alzate, C. A., Roman-Valencia, C. & Taphorn, D. C. (2013) Una nueva especie de Hyphessobrycon (Characiformes: Characidae) de la cuenca del rio Telembi, vertiente sur del Pacifico, Colombia. Revista de Biologia Tropical, 61 (1), 181 - 192. http: // doi. org / 10.15517 / RBT. V 61 I 1.10944","Garcia-Alzate, C. A., Roman-Valencia, C., Taphorn, D. C. & Villa-Navarro, F. A. (2015) Hyphessobrycon natagaima (Characiformes: Characidae) a new species from Colombia, with a key to the Magdalena Basin Hyphessobrycon species. Caldasia, 37 (1), 221 - 232. https: // doi. org / 10.15446 / caldasia. v 37 n 1.51228","DoNascimiento, C., Herrera-Colazos, E. E., Herrera-R, G. A., Ortega-Lara, A., Villa-Navarro, F. A., Usma Oviedo, J. S. & Maldonado-Ocampo, J. A. (2017) Checklist of the freshwater fishes of Colombia: a Darwin Core alternative to the updating problem. ZooKeys, 708, 25 - 138. http: // doi. org / 10.3897 / zookeys. 708.13897","Eigenmann, C. H. (1918) The American Characidae [Part II]. Memories of the Museum of Comparative Zoology, 43, 103 - 208, pls. 9 - 11, 13, 17 - 29, 33, 78 - 80, 93.","Eigenmann, C. H. (1920 a) South America West of Maracaibo, Orinoco, Amazon, and Titicaca basins, and the horizontal distribution of its fresh-water fishes. Indiana University Studies, (45), 1 - 24.","Eigenmann, C. H. (1920 b) The fresh-water fishes of Panama East of longitude 80 º. Indiana University Studies, (175), 3 - 19.","Eigenmann, C. H. (1920 c) The fishes of the rivers draining the Western slope of the cordillera occidental of Colombia, rios Atrato, San Juan, Dagua, and Patia. Indiana University Studies, (1181), 1 - 19.","Eigenmann, C. H. (1920 d) The Magdalena basin and the horizontal and vertical distribution of its fishes. Indiana University Studies, (177), 21 - 34.","Dahl, G. (1971) Los peces del norte de Colombia. Bogota: Inderena.","Weitzman, S. H. & Palmer, L. (1997) A new species of Hyphessobrycon (Teleostei: Characidae) from the Neblina region of Venezuela and Brazil, with comments on the putative 'rosy tetra clade'. Ichthyological Exploration of Freshwaters, 7 (3), 209 - 242."]}

Hyphessobrycon daguae Eigenmann, 1922 (Figs. 11���14, Tab. 4) Hyphessobrycon panamensis daguae Eigenmann, 1922: 141 [original description, type locality: ���Dagua and Pat��a basins���, Colombia]���. Eigenmann & Myers, 1929: 534���535 [comments on the taxon]���. Ibarra & Stewart, 1987: 48 [type catalog]���. Zarske & G��ry, 2002: 26 [possible junior synonym of Hyphessobrycon condotensis]���.Lima et al., 2003: 139 [listed; as a synonym of Hyphessobrycon panamensis]. Hyphessobrycon panamensis (not Durbin) ���. Eigenmann, 1918: 186 [in part; specimens from r��o Pat��a]���. Eigenmann, 1920a: 17 [in part; specimens from r��o Pat��a]���. Barriga, 1994: 73 [list of species, northwest of Ecuador]���. Eigenmann, 1922: 141 [in part; specimens from Colombia: Raspadura, and Tambo]���. Weitzman & Palmer, 1997: 228 [in part: specimens from r��o Esmeraldas, r��o Santiago, r��o Bogot��, Estero Sabalera, Ecuador; comments on color in life and distribution]���.Lima et al., 2003: 139 [as a synonym of H. panamensis]. Hyphessobrycon ecuadoriensis (not Eigenmann & Henn) ���. Barriga, 2012: 107 [list of species, Ecuador]���. Garc��a-Alzate et al., 2013: 184 [in part: MEPN 85-117]. Hyphessobrycon chocoensis Garcia-Alzate, Rom��n-Valencia & Taphorn, 2013: 183 [original description, type locality: ��� Colombia, Nari��o, Barbacoas, Quebrada la Tundera afluente del r��o Quigualp�����]���. Maldonado-Ocampo et al., 2012: 115 [Colombia, Choc��; photograph; short description; distribution] ���. DoNascimiento et al., 2017: 49 [listed, Colombia]. NEW SYNONYM. Hyphessobrycon sp. nov. Jim��nez-Prado et al., 2015: 139 [brief description; biological data; distribution]. Diagnosis. Hyphessobrycon daguae differs from all congeners, except from H. compressus, H. diancistrus, H. otrynus, and those belonging to the H. panamensis species-group, by the presence of one large bony hook per fin ray, antero-dorsally oriented and surrounded by a fleshy area, on last unbranched and first branched anal-fin rays of mature males (vs. absence of hooks, or presence of small and numerous hooks by ray). Hyphessobrycon daguae can be distinguished from H. compressus by the absence of a black blotch on the dorsal fin (vs. presence), by the presence of predorsal series (vs. absence), and by presenting 32���35 scales in longitudinal series (vs. 41���48). It can be distinguished from H. diancistrus and H. otrynus by presenting the caudal fin hyaline (vs. dark blotches on caudalfin lobes in H. diancistrus; distal third of caudal fin darkened in H. otrynus). It differs from the remaining species of the H. panamensis group by presenting only one diffuse humeral blotch (vs. two humeral blotches, the first one well defined), and by presenting the lateral line frequently disrupted (vs. incompletely pored, or rarely disrupted in H. condotensis). It can be further distinguished from H. columbianus by the absence of a silvery-blue iridescent coloration on dorsal region of flank in life (vs. presence), and by the lower body depth (39.8���43.0% SL vs. 46.0���50.3% SL). It can be distinguished from H. savagei by presenting one diffuse humeral blotch, extending one longitudinal scale row below the lateral line (vs. two humeral blotches equally pigmented and always well defined, extending two to four longitudinal scale rows below the lateral line). It also differs from H. panamensis by a higher number of branched anal-fin rays (23���27, mode = 24 vs. 19���23, modes = 21 and 22). It can be additionally distinguished from H. bussingi and H. condotensis by presenting 7���8 (mode = 7) scale rows between lateral line and dorsal-fin insertion (vs. 6���7, mode = 6�� in H. bussingi; and 5���7, mode = 6 in H. condotensis). Description. Morphometric data in Table 4. Body compressed, greatest depth at vertical through dorsal-fin origin. Dorsal profile of body convex from upper lip to vertical through anterior nostril; straight or slightly concave from this point to distal tip of supraoccipital process. Slightly convex from tip of supraoccipital process to dorsal-fin origin; straight or slightly convex from this point to adipose-fin origin; and slightly concave along caudal peduncle. Ventral profile of body convex from lower lip to pelvic-fin origin; straight or slightly convex from this point to analfin origin; straight or slightly convex along anal-fin base; and slightly concave along caudal peduncle. Jaws equal, mouth terminal. Premaxillary teeth in two rows; outer row with 2(2), 3(4), 4(11), or 5(2) tricuspid teeth; inner row with 4(1) or 5(25) tri- to pentacuspid teeth. Posterior margin of maxilla slightly posterior to vertical through anterior margin of pupil. Maxilla with 1(2), 2(18), or 3(6) conical to tricuspid teeth. Dentary with 4(10), 5(14), or 6(4) large penta and tricuspid teeth, gradually decreasing in size, followed by up to 11 conical and abruptly smaller teeth. Central cusp longer than lateral ones. Scales cycloid. Lateral line incompletely pored with 8(1), 9(1), 10(1), 11(1), 13(1), 14*(1), 16(2), 21(1), 25(1), 31(1), disrupted with total of 15(1), 16(1), 18(1), 20(1), 21(1), 23(1), 26(1), 27(1), 28(1), 29(1), 30(1), 33(2), or completely pored with 35(1) perforate scales. Longitudinal series, including perforate scales, with 32(7), 33(5), 34(3), 35(6). Scale rows between dorsal-fin origin and lateral line 7(19) or 8(2). Scale rows between lateral line and pelvic-fin insertion 5(3), 5��(2), or 6(16). Predorsal series with 10(15) or 11(8) scales. Single row of ten scales covering base of anteriormost anal-fin rays. Circumpeduncular scales 13(6) or 14(16). Dorsal-fin rays ii,7,ii(4), 8,i(9), 9(9), or 10(1). Adipose fin present. Pectoral-fin rays i,10(2), 11(10), or 12(10), distal tip of rays surpassing pelvic-fin origin. Pelvic-fin rays i,7(21), distal tip surpassing anal-fin origin. Anal-fin rays iii(1) or iv(26), 23(5), 24(11), 25*(4), 26(6), or 27(1). Anal fin of mature males with large bony hooks. Caudal fin bifurcate, lobes approximately of same size, i,9/8,i rays. First gill arch with 6(3), 7(16), or 8(2) rakers on epibranchial, 1(15) on intermediate cartilage, 11(4), 12(11), or 13(5) on cerato+hipobranchial. Branchiostegal rays 4. Color in alcohol. Background coloration beige. Dorsal region of head and middorsal region of body with dense concentration of melanophores. Region above pupil darkened. Infraorbitals and opercle beige, with few scattered melanophores. Opercle with some guanine pigmentation. Lower portion of maxilla and gular region with few scattered melanophores, slightly more concentrated on upper half. Single humeral blotch, diffuse and vertically elongat- ed, with scattered melanophores equally distributed, extending three longitudinal scale rows above and one below lateral line. Body sides with melanophores evenly distributed, except on anteroventral half, with few melanophores. Posterior margin of scales on superior third of flanks with concentration of melanophores. Brownish-gray longitudinal midlateral stripe, from humeral blotch to caudal peduncle, more conspicuous on posterior region. Fins hyaline, Color in life. (Based in Jimen��z-Prado et al. 2015: 141, as Hyphessobrycon sp. nov.). Background color silvery to beige. Dark markings as in preserved specimens. Dorsal region of body brownish-gray. Upper half of head brownish-gray; lower half of head silvery to beige. Humeral blotch absent or inconspicuous. Brownish-gray longitudinal midlateral stripe. Abdominal region whitish-yellow. Fins hyaline; rosy or reddish in living specimens. Distribution. Hyphessobrycon daguae is known from the r��o Dagua, r��o Pat��a and r��o Telemb�� basins in Colombia, and r��o Mira (at border between Colombia and Ecuador), r��o Cayapas, r��o Mataje, and r��o Santiago basins, in Ecuador (Figs. 13, and 21). Sexual dimorphism. Mature males with one pair of large bony hooks per fin ray, antero-dorsally oriented and surrounded by a fleshy area, from the last unbranched (on the fourth segment) to the third branched (varying on fourth to seventh segments) anal-fin ray of mature males. Very small hooks, difficult to discern, can be present on distal segments of most anterior anal-fin rays. Dorsal-fin length of males much longer than in females. Ecology and habitat. Hyphessobrycon daguae occur in watercourses with slow flowing, clear water, and abundant riparian vegetation (Jim��nez-Prado et al. 2015, as Hyphessobrycon sp. nov.). Remarks. Eigenmann (1922) described Hyphessobrycon daguae as a subspecies, H. panamensis daguae. In the original description, S.F. Hildebrand highlighted that its lateral line was ���frequently almost completely pored���. Despite that, subsequently to its description, H. panamensis daguae has been considered as a junior synonym of H. condotensis or of H. panamensis. However, H. condotensis and H. panamensis present the lateral line typically incompletely pored (except for one specimen of H. condotensis, ICNMHN 205, with lateral line disrupted; and one specimen of H. panamensis, UCR 1463006, with lateral line completely pored). Additionally, H. daguae can be distinguished from these species by presenting a single diffuse humeral blotch (vs. two humeral blotches, the first one always well defined, even if inconspicuous). Hyphessobrycon daguae presents polymorphism on the lateral line (which can be incompletely, disrupted, or completely pored) similar to that found in some other Characidae, such as H. balbus Myers (Burger et al., 2019), and Hemigrammus ataktos Marinho, Birindelli & Dagosta (Marinho et al., 2014). This condition, as well as other characters, are similar to those presented in the description of Hyphessobrycon chocoensis, a species described from the r��o Telemb�� basin, which is a tributary of the r��o Pat��a (type locality of H. daguae). Furthermore, two syntypes of H. daguae (FMNH 56527 and FMNH 56528) are exactly from the same municipality and drainage of the type series of H. chocoensis. Therefore, as no differences between these nominal species were found, H. daguae is herein revalidated and H. chocoensis is considered as its junior synonym. The type series of Hyphessobrycon panamensis daguae is composed by nine syntypes (FMNH 56156, FMNH 56526, FMNH 56527, FMNH 56528, FMNH 56529, and FMNH 58316). The lectotype herein designated is the 26.0 mm SL specimen of FMNH 58316 (Fig. 11a), and the remaining syntypes become thus paralectoytpes, as ruled by the article 74 of ICZN (1999). Another issue involving Hyphessobrycon daguae concerns the specimens from the province of Esmeraldas, Northern Ecuador. Weitzman & Palmer (1997: 228-229) identified the specimens from the r��o Esmeraldas as H. panamensis, while Barriga (2012) recorded H. ecuadoriensis from the r��o Santiago (a river drainage very close to r��o Esmeraldas). On the other hand, Garc��a-Alzate et al. (2013) recorded H. ecuadoriensis from the r��o Mira (at the border between Colombia and Ecuador) to the drainages of Southern Ecuador. However, Jim��nez-Prado et al. (2015) restricted the distribution of H. ecuadoriensis to the r��o Guayas basin (southern Ecuador), and identified the specimens from the r��o Mataje and r��o Santiago basins as ��� Hyphessobrycon sp. nov.���, distinguishing it from H. daguae (as H. chocoensis) by the presence of seven perforated scales on the lateral line (vs. 10 or more). Nonetheless, the specimens from the r��o Bravo (MEPN 3701) and from the r��o Santiago basins (ZUEC 11330) can present the lateral line incompletely pored or disrupted (Fig. 14). This emphasizes the polymorphic condition of the character for those populations, and, consequently, the count of perforated scales in ��� Hyphessobrycon sp. nov.��� is herein considered to be best interpreted as a population variation within H. daguae. Therefore, the distribution of Hyphessobrycon daguae is extended herein to the r��o Mira, r��o Cayapas, r��o Mataje, and r��o Santiago basins, in Ecuador. Material examined. 34 specimens (15.6���43.4 mm SL). Type material. FMNH 58316, lectotype of Hyphessobrycon daguae (by present designation), 26.0 mm SL: Colombia, r��o Pat��a basin; A. W. Henn, 1913. FMNH 144330, paralectotype of Hyphessobrycon daguae, 28.0 mm SL, same data as lectotype. IUQ 3035, holotype of Hyphessobrycon chocoensis, 35.3 mm SL: Colombia, Nari��o, Barbacoas, creek la Tundera, tributary of the r��o Quigualp��, 01��39���39������N 78��09���04������W; C. Rom��n-Valencia, C. Garc��a-Alzate, R. Ruiz-C, V. Tovar Sr., V. Tovar Jr. & H. Cortes, 16 Jul 2008; IUQ 2274, paratypes of Hyphessobrycon chocoensis, 6, 15.6���40.6 mm SL: Colombia, Nari��o, Barba- coas, upper r��o Yamund��, tributary of the r��o Telemb��, 01��39���39������N 78��09���04������W; C. Rom��n-Valencia, C. Garc��a- Alzate, R. Ruiz-C, V. Tovar Sr., V. Tovar Jr. & H. Cortes, 16 Jul 2008; IUQ 2275, paratypes of Hyphessobrycon chocoensis, 7, 24.9���33.4 mm SL, same data as holotype. Non-type material. Colombia, Nari��o, r��o Telembi basin: IUQ 2311, 3, 21.5���34.4 mm SL, creek on Hacienda, tributary of the r��o Telembi, 01��30���N 77��58���W, 16 Jul 2008. Ecuador, Esmeraldas: MEPN 3697, 4, 34.1���43.4 mm SL, tributary of the r��o Bravo, 8 km above r��o Bravo falls, r��o Cayapas, 00��38���22������N 78��55���24������W; R. Barriga, C. Mina & W. Nazareno, 18 Jul 1985; MEPN 3701, 6, 35.0��� 43.3 mm SL, same data as MEPN 3697; ZUEC 11330, 5, 30.5 ��� 27.8 mm SL, Estero la Chiquita, tributary of the r��o Santiago, 1��13���50���N, 78��45���17���W; R. Barriga & C. Mina, 21 Oct 1985., Published as part of Ota, Renata R., Carvalho, Fernando R. & Pavanelli, Carla S., 2020, Taxonomic review of the Hyphessobrycon panamensis species-group (Characiformes: Characidae), pp. 401-436 in Zootaxa 4751 (3) on pages 415-421, DOI: 10.11646/zootaxa.4751.3.1, http://zenodo.org/record/3714610, {"references":["Eigenmann, C. H. (1922) The fishes of Western South America [Part I]: The fresh-water fishes of Northwestern South America, includind Colombia, Panama, and the Pacific slopes of Ecuador and Peru, together with and appendix upon the fishes of the rio Meta in Colombia. Memoirs of the Carnegie Museum, 9 (1), 1 - 346.","Eigenmann, C. H. & Myers, G. S. (1929) The American Characidae [Part V]. Memories of the Museum of Comparative Zoology, 43, 429 - 558, pls. 57, 63, 70 - 74, 81 - 83, 94.","Ibarra, M. & Stewart, D. J. (1987) Catalogue of type specimens of recent fishes in Field Museum of Natural History. Fieldiana, Zoology, 35, 1 - 112.","Zarske, A. & Gery, J. (2002) Der Blaurote Kolumbien-Salmler. Hyphessobrycon columbianus n. sp. - ein neuer Salmler (Teleostei; Characiformes, Characidae) aus dem kolumbianischen Darien. Das Aquarium, 36 (391), 22 - 30.","Eigenmann, C. H. (1918) The American Characidae [Part II]. Memories of the Museum of Comparative Zoology, 43, 103 - 208, pls. 9 - 11, 13, 17 - 29, 33, 78 - 80, 93.","Eigenmann, C. H. (1920 a) South America West of Maracaibo, Orinoco, Amazon, and Titicaca basins, and the horizontal distribution of its fresh-water fishes. Indiana University Studies, (45), 1 - 24.","Barriga, R. (1994) Peces del Noroeste del Ecuador. Politecnica, 19 (2), 43 - 154.","Weitzman, S. H. & Palmer, L. (1997) A new species of Hyphessobrycon (Teleostei: Characidae) from the Neblina region of Venezuela and Brazil, with comments on the putative 'rosy tetra clade'. Ichthyological Exploration of Freshwaters, 7 (3), 209 - 242.","Barriga, R. (2012) Lista de peces de agua dulce e intermareales del Ecuador. Politecnica, 30 (3), 83 - 119.","Garcia-Alzate, C. A., Roman-Valencia, C. & Taphorn, D. C. (2013) Una nueva especie de Hyphessobrycon (Characiformes: Characidae) de la cuenca del rio Telembi, vertiente sur del Pacifico, Colombia. Revista de Biologia Tropical, 61 (1), 181 - 192. http: // doi. org / 10.15517 / RBT. V 61 I 1.10944","Maldonado-Ocampo, J. A., Usma, J. S., Villa-Navarro, F. A., Ortega-Lara, A., Prada-Pedreros, S., Jimenez, L. F., Karamillo-Villa, U., Arango, A., Rivas, T. & Sanchez, G. C. (2012) Peces dulceacuicolas del Choco biogeografico de Colombia. WWF Colombia, IAvH, AU-NAP, Pontificia Universidad Javeriana, Bogota (DC).","DoNascimiento, C., Herrera-Colazos, E. E., Herrera-R, G. A., Ortega-Lara, A., Villa-Navarro, F. A., Usma Oviedo, J. S. & Maldonado-Ocampo, J. A. (2017) Checklist of the freshwater fishes of Colombia: a Darwin Core alternative to the updating problem. ZooKeys, 708, 25 - 138. http: // doi. org / 10.3897 / zookeys. 708.13897","Jimenez-Prado, P., Aguirre, W., Laaz-Moncayo, E., Navarrete-Amaya, R., Nugra-Salazar, F., Rebolledo-Monsalve, E., Zarate- Hugo, E., Torres-Noboa, A. & Valdiviezo-Rivera, J. (2015) Guia de peces para aguas continentales en la vertiente occidental del Ecuador. Pontificia Universidad Catolica del Ecuador Sede Esmeraldas (PUCESE), Universidad del Azuay (UDA) y Museo Ecuatoriano de Ciencias Naturales (MECN) del Instituto Nacional de Biodiversidad, Esmeraldas.","Burger, R., Carvalho, F. R. & Zanatta, A. M. (2019) A new species of Astyanax Baird & Girard (Characiformes: Characidae) from western Chapada Diamantina, Bahia, Brazil. Zootaxa, 4604, 369 - 380. http: // doi. org / 10.11646 / zootaxa. 4604.2.9","Marinho, M. M. F., Dagosta, F. C. P. & Birindelli, J. L. O. (2014) Hemigrammus ataktos: a new species from the rio Tocantins basin, central Brazil (Characiformes: Characidae). Neotropical Ichthyology, 12 (2): 257 - 264. http: // doi. org / 10.1590 / 1982 - 0224 - 20130091"]}

Hyphessobrycon panamensis Durbin, 1908 (Figs. 15���18, Tab. 5) Hyphessobrycon panamensis Durbin, in Eigenmann, 1908: 101 [original description, type locality: ���Boqueron River, Panama ���]���. Eigenmann, 1910: 436 [Panama; listed]���. Meek & Hildebrand, 1916: 287 [Panama, r��o Chagres basin; synonymic list; description; distribution]���. Eigenmann, 1918: 186, plate 26 (fig. 1), plate 33 (fig. 4) [in part; specimens from Panama, r��o Boqueron basin; synonymic list; description; distribution]���. Eigenmann, 1920a: 17 [in part; specimens from Panama, r��o Chagres basin; distribution]���. Eigenmann, 1920b: 15 [in part; specimens from Panama, r��o Chagres; distribution]���. Eigenmann, 1922: 141 [synonymic list; in part; specimens from Panama, r��o Chagres; distribution]���. Jordan et al., 1930 [Panama, r��o Chagres basin; synonymic list; distribution]���. Hildebrand, 1938: 269 [Panama, r��o Chagres basin; short description]���. Vari & Howe, 1991: 27 [type catalog]���. Weitzman & Palmer, 1997: 227���229 [in part; specimens from Panama: canal Zone, r��o Chagres, r��o Tocu��, ���Largateria creek���, upper Trinidad, Reservois Creek; comments on color in life and distribution]���. Zarske & G��ry, 2002: 22-30 [diagnosis from H. columbianus; comments on cryptic species; diagnosis in key, photo]���.Lima et al., 2003: 139 [listed; distribution]���. Maldonado-Ocampo et al., 2008: 169 [listed, Colombia; Pacific slope rivers; distribution]���. Garc��a-Alzate et al., 2010: 55 [recorded from Colombia; comparison with H. sebastiani]���. Garc��a-Alzate et al., 2013: 182 [restriction of the distribution to Panama and Costa Rica; in part; specimens from Panama: r��o Boqueron, and zona del Canal del Panama]. Hemigrammus minutus Meek & Hildebrand, 1912: 67 [original description, type locality: ���r��o Agua Clara, Canal Zone, Panama ���]���. Eigenmann, 1913: 31 [synonymization with Hyphessobrycon panamensis]���. Meek & Hildebrand, 1916: 287 [as a synonym of H. panamensis]���. Eigenmann, 1918: 186 [as a synonym of H. panamensis]���. Ibarra & Stewart, 1987: 45 [type catalog]���. Weitzman & Palmer, 1997: 227 [as a synonym of H. panamensis]���. Zarske & G��ry, 2002: 26���28 [as a synonym H. panamensis;]���.Lima et al., 2003: 139 [listed; as a synonym of H. panamensis]. Diagnosis. Hyphessobrycon panamensis differs from all congeners, except from H. compressus, H. diancistrus, H. otrynus, and those belonging to the H. panamensis species-group, by the presence of one large bony hook per fin ray, antero-dorsally oriented and surrounded by a fleshy area, on last unbranched and first branched anal-fin rays of mature males (vs. absence of hooks, or presence of small and numerous hooks on anteriormost anal-fin rays). Hyphessobrycon panamensis can be distinguished from H. compressus by the absence of a dark blotch on the dorsal fin (vs. presence), by the presence of predorsal series (vs. absence), and by presenting 33���35 scales in longitudinal series (vs. 41���48). It can be distinguished from H. diancistrus and H. otrynus by presenting the caudal fin hyaline (vs. presence of dark blotches on caudal-fin lobes in H. diancistrus; distal third of caudal fin darkened in H. otrynus). It differs from H. columbianus by the absence of a silvery-blue iridescent coloration on dorsal region of flank in life (vs. presence), and by the lower body depth (36.1���45.7% SL vs. 46.0���50.3% SL in H. columbianus). It can be distinguished from H. savagei by presenting the second humeral blotch less conspicuous than the first, extending one or two longitudinal scale rows below lateral line, with posterior margin as a concentration of melanophores dispersing towards longitudinal midlateral stripe (vs. humeral blotches equally pigmented, extending two to four longitudinal scale rows below lateral line, with contours and margins always well-defined). Hyphessobrycon panamensis differs from H. bussingi, H. condotensis and H. daguae by the lower number of branched anal-fin rays (19���23, modes = 21 and 22; vs. 23���27, rarely 22, mode = 24 in H. condotensis and H. daguae, and mode = 25 in H. bussingi). It can be further distinguished from H. bussingi and H. condotensis by presenting 6, rarely 5�� (mode = 6), scale rows between lateral line and pelvic-fin insertion (vs. 4���5��, mode = 5 in H. bussingi; and 4�����6, mode = 5 in H. condotensis). It also differs from H. condotensis, by presenting 7���8 (mode = 7) scale rows between lateral line and dorsal-fin insertion (vs. 5���7, mode = 6). Additionally, it can be distinguished from H. daguae by presenting a well-defined first humeral blotch (vs. only one diffuse humeral blotch). Description. Morphometric data in Table 5. Body compressed, greatest depth at vertical through dorsal-fin origin. Dorsal profile of body convex from upper lip to vertical through anterior nostril; straight or slightly concave from this point to distal tip of supraoccipital process. Slightly convex from tip of supraoccipital process to dorsal-fin origin; straight or slightly convex from this point to adipose-fin origin; and straight along caudal peduncle. Ventral profile of body convex from lower lip to pelvic-fin origin; straight or slightly convex from this point to anal-fin origin; straight along anal-fin base; and straight or slightly concave along caudal peduncle. Jaws equal, mouth terminal. Premaxillary teeth in two rows; outer row with 2(1), 3*(9), 4(2), or 5(1) tri- to pentacuspid teeth; inner row with 5*(15) tri- to pentacuspid teeth. Posterior margin of maxilla at vertical through anterior margin of pupil. Maxilla with 2(6), 3*(6), 4(1), or 5(2) tri- to pentacuspid teeth. Dentary with 4*(8), 5(6) large penta- and tricuspid teeth, gradually decreasing in size, followed by conical and abruptly smaller teeth. Central cusp more developed than lateral ones. Scales cycloid. Lateral line incompletely pored with 7(1), 8(2), 9(2), 10(6), 11*(6), 12(4), 13(2), 15(1), 16(1), or completely pored with 33(1) perforate scales. Longitudinal series, including perforate scales, with 33*(6), 34(3), or 35(3). Scale rows between dorsal-fin origin and lateral line 7(19), 7��*(1), or 8(3). Scale rows between lateral line and pelvic-fin insertion 5��*(2) or 6(20). Predorsal series with 11*(4), 12(5), or 13(2) scales. Single row of six scales covering base of anteriormost anal-fin rays. Circumpeduncular scales 13(1), 14*(7), or 15(2). Dorsal-fin rays ii,8(1) or ii,9*(14). First dorsal-fin pterygiophore located between neural spines of ninth and 10 th (3) vertebrae. Adipose fin present. Pectoral-fin rays i,10*(3), 11(7), or 12(2), distal tip of rays reaching pelvic-fin origin. Pelvic-fin rays i,7*(13), distal tip reaching anal-fin origin. Anal-fin rays iii*(17), iv(6), 19(1), 20(2), 21(8), 22*(8), or 23(4). Anal fin of mature males with large bony hooks. Caudal fin bifurcate, lobes approximately of same size, i,9/8,i(1) rays. First gill arch with 7(7) or 8(2) rakers on epibranchial, 1(9) on intermediate cartilage, 10(1), 11(6), 12(1), or 13(1) on cerato+hipobranchial. Branchiostegal rays 4. Total vertebrae 33*(3), supraneurals 4*(2) or 5(1) (Fig. 17). Color in alcohol. Background coloration beige to light brown. Dorsal region of head and middorsal region of body with dense concentration of melanophores. Region above pupil darkened. Infraorbitals beige, with few scattered melanophores. Opercle retaining guanine pigmentation, without melanophores. Inferior portion of maxilla and gular region with few scattered melanophores, slightly more concentrated on anterior half. Two dark-brown humeral blotches, vertically elongated, separated by light-beige area. First humeral blotch conspicuous, with melanophores equally distributed, extending four to five longitudinal scale rows above and two below lateral line. Second humeral blotch usually inconspicuous, extending four to five longitudinal scale rows above and one below lateral line; posterior margin with concentration of melanophores concentrated along longitudinal midlateral stripe. Body sides with melanophores regularly distributed, except on antero-ventral half, with few or without melanophores. Posterior margin of scales on superior third of body sides with concentration of melanophores. Brownish to silvery longitudinal midlateral stripe, from second humeral blotch to caudal peduncle, more conspicuous on posterior region. Fins hyaline, with slight concentration of melanophores on distal half of dorsal fin, distal third of anal fin, and median caudal-fin rays. Abdominal region, from pectoral-fin origin to pelvic-fin end, without melanophores (Fig. 18). Distribution. Hyphessobrycon panamensis is known from the r��o Bejuco, r��o Boquer��n, r��o Calov��bora, r��o Chagres and r��o Lllano Succio basins, and in the Canal Zone, in Panama (both Atlantic and Pacific versants) (Figs. 4, 8 and 21). Sexual dimorphism. Mature males with one large bony hook per fin ray, anterodorsally oriented and surround- ed by fleshy area, from the last unbranched (on fourth segment) to the third branched (varying on fifth to seventh segments) anal-fin rays of mature males (Figs. 5b and 6b). Very small hooks, difficult to discern, can be present on distal segments of most anterior anal-fin rays. Dorsal-fin length of males slightly longer than in females. Ecology and habitat. Hyphessobrycon panamensis feeds on aquatic and terrestrial insects. The oocytes diameter varies between 0.7 to 0.8 mm (Kramer, 1978; Kramer & Bryant, 1995). Remarks. Eigenmann (1913) proposed Hemigrammus minutus Meek & Hildebrand as a junior synonym of Hyphessobrycon panamensis. The examination of the paratypes of He. minutus conducted during the present study showed that the purported diagnostic characters from the latter taxon overlap with H. panamensis and, therefore, the proposed synonymy is herein confirmed. Hyphessobrycon panamensis has its diagnosis from similar species redefined herein based mainly in the low number of branched anal-fin rays. The geographical distribution of the species is restricted herein to the central provinces of Panama, draining both Atlantic and Pacific versants. There is no record indicating the syntopy with other species of the group. In the literature, Hyphessobrycon panamensis has been recorded in Costa Rica (e.g. Bussing, 1987, 1998) and Colombia (e.g. Eigenmann, 1920a, b, c, d; Dahl, 1971; Weitzman & Palmer, 1997; Zarske & G��ry, 2002; Garc��a-Alzate et al., 2010). However, specimens identified as H. panamensis from Colombia belong either to H. columbianus, H. condotensis or H. daguae (the latter species herein revalidated and considered as senior synonym of H. chocoensis; see remarks on this species, above). In addition, specimens identified as H. panamensis from the province of San Blas, southeast Panama actually correspond to H. columbianus. On the other hand, specimens previously identified as Hyphessobrycon panamensis from Costa Rica and in the province of Bocas del Toro, Panama (at the border with Costa Rica), belong rather to H. bussingi, new species, described herein. The type series of Hyphessobrycon panamensis is composed of six syntypes (MCZ 20688 and USNM 120416). The lectotype herein designated is a male specimen with large hooks on the anal-fin ray (USNM 120416, fig. 15a), and all the remaining specimens consequently become paralectotypes, as ruled by the article 74 of ICZN (1999). Material examined. 54 specimens (20.6���48.6 mm SL). All from Panama. Type material. USNM 120416, lectotype of Hyphessobrycon panamensis (by present designation), 25.4 mm SL (rd): r��o Boqueron, Atlantic slope; Hassler Expedition, 31 Jul 1872. USNM 443817, paralectotype of Hyphessobrycon panamensis, 25.7 mm SL (rd), same data as lectotype. MCZ 20688, paralectotypes of Hyphessobrycon panamensis, 5, 20.6���23.5 mm SL (dry specimens), same data as lectotype; FMNH 12770, paratypes of Hemigrammus minutus, 2, 25.3���30.5 mm SL: Panama, Cana Zone, Agua Clara; S. F. Hildebrand & S. E. Meek, 10 Mar 1911; FMNH 12771, paratype of Hemigrammus minutus, 35.9 mm SL: Panama, Canal Zone, r��o Mandingo, Bas Obispo; S.F. Hildebrand & S.E. Meek, 02 Feb 1911. USNM 78628, paratypes of Hemigrammus minutus, 31 (20, 22.9���31.3 mm SL): Canal Zone, r��o Agua Clara; S.E. Meek & S.F. Hildebrand, 10 Mar 1911. Non-types. Bocas del Toro, r��o Calov��bora basin: UCR 1463006, 2 (1, 34.6 mm SL), r��o Calou��bora, next to mouth, 08��33��N 81��11��W; T. Thomson, 01 Apr 1983. C��lon, r��o Llano Sucio basin: AMNH 37802, 24 (4, 25.5���31.4 mm SL): Canal Zone, quebrada Juan Grande near Gamboa, 09��07���N 79��43���W; P.C. Rosen et al., 04 Jan 1978. LBP 2759, 7, 26.6���35.0 mm SL (1 rd, 34.8 mm SL): Santa Rita de Arriba, r��o Llano Sucio, 09��19���26.2������N 79��46���08.2������W; C. Oliveira, R. G. Reina, C. Veja & S. Perez, 14 Jul 2005. Veraguas, r��o Bejuco basin: MCP 39195, 5, 36.0 ��� 48.6 mm CP: r��o Bejuco, 08��48���N 81��01���W; P. Rham, 12 Aug 1998. West Panama, Canal Zone: ANSP 99866, 16 (5, 31.9���35.5 mm SL), creek 2 mi E of Nuevo Emperador road, 9��02���N 79��47���W; H.G. Loftin & E.L. Tyson, 17 Aug 1962. ANSP 146800, 14 (5, 24.7���30.5 mm SL), brook 1.1 km E of Nuevo Emperador on Nuevo Emperador-Arraijan road, 9��02���N 79��47���W; D. Fromm & P. Fromm, 13 Mar 1981., Published as part of Ota, Renata R., Carvalho, Fernando R. & Pavanelli, Carla S., 2020, Taxonomic review of the Hyphessobrycon panamensis species-group (Characiformes: Characidae), pp. 401-436 in Zootaxa 4751 (3) on pages 422-426, DOI: 10.11646/zootaxa.4751.3.1, http://zenodo.org/record/3714610, {"references":["Eigenmann, C. H. (1908) Zoological Results of the Thayer Brazilian expedition: Preliminary descriptions of new genera and species of tetragonopterid characins. Bulletin of the Museum of Comparative Zoology, 52 (6), 91 - 106.","Eigenmann, C. H. (1910) Catalogue of the fresh-water fishes of Tropical and South temperate America. In: Scott, WB (Ed.), Reports of the Princeton University expeditions to Patagonia 1896 - 1899. Princeton University, pp. 375 - 511, vol. 3, Zoology, pt. 4.","Meek, S. E. & Hildebrand, S. F. (1916) The fishes of the fresh waters of Panama. The Field Museum of Natural History, Zoological Series, 10 (15), 1 - 374.","Eigenmann, C. H. (1918) The American Characidae [Part II]. Memories of the Museum of Comparative Zoology, 43, 103 - 208, pls. 9 - 11, 13, 17 - 29, 33, 78 - 80, 93.","Eigenmann, C. H. (1920 a) South America West of Maracaibo, Orinoco, Amazon, and Titicaca basins, and the horizontal distribution of its fresh-water fishes. Indiana University Studies, (45), 1 - 24.","Eigenmann, C. H. (1920 b) The fresh-water fishes of Panama East of longitude 80 º. Indiana University Studies, (175), 3 - 19.","Eigenmann, C. H. (1922) The fishes of Western South America [Part I]: The fresh-water fishes of Northwestern South America, includind Colombia, Panama, and the Pacific slopes of Ecuador and Peru, together with and appendix upon the fishes of the rio Meta in Colombia. Memoirs of the Carnegie Museum, 9 (1), 1 - 346.","Jordan, D. S., Warren-Evermann, B. & Clark, H. W. (1930) Check list of the fishes and fishlike vertebrates of North and Middle America north of the northern boundary of Venezuela and Colombia [Part II]. Report of the United States Commissioner of Fisheries, 1 - 670.","Hildebrand, S. F. (1938) A new catalog of the fresh-water fishes of Panama. The Field Museum of Natural History, Zoological Series, 22 (4), 219 - 359.","Vari, R. P. & Howe, J. C. (1991) Catalog of type specimens of recent fishes in the National Museum of Natural History, Smithsonian Institution [Part I]: Characiformes (Teleostei, Ostariophysi). Smithsonian Contributions to Zoology, 517, 1 - 52.","Weitzman, S. H. & Palmer, L. (1997) A new species of Hyphessobrycon (Teleostei: Characidae) from the Neblina region of Venezuela and Brazil, with comments on the putative 'rosy tetra clade'. Ichthyological Exploration of Freshwaters, 7 (3), 209 - 242.","Zarske, A. & Gery, J. (2002) Der Blaurote Kolumbien-Salmler. Hyphessobrycon columbianus n. sp. - ein neuer Salmler (Teleostei; Characiformes, Characidae) aus dem kolumbianischen Darien. Das Aquarium, 36 (391), 22 - 30.","Maldonado-Ocampo, J. A., Vari, R. P. & Usma, J. S. (2008) Checklist of the freshwater fishes of Colombia. Biota Colombiana, 9 (2), 143 - 237.","Garcia-Alzate, C. A., Roman-Valencia, C. & Taphorn, D. C. (2010) A new species of Hyphessobrycon (Teleostei: Characiformes: Characidae) from the San Juan River drainage, Pacific versant of Colombia. Zootaxa, 2349, 55 - 65. http: // doi. org / 10.11646 / zootaxa. 2349.1.4","Garcia-Alzate, C. A., Roman-Valencia, C. & Taphorn, D. C. (2013) Una nueva especie de Hyphessobrycon (Characiformes: Characidae) de la cuenca del rio Telembi, vertiente sur del Pacifico, Colombia. Revista de Biologia Tropical, 61 (1), 181 - 192. http: // doi. org / 10.15517 / RBT. V 61 I 1.10944","Meek, S. E. & Hildebrand, S. F. (1912) Descriptions of new fishes from Panama. The Field Museum of Natural History, Zoological Series, 10 (6), 67 - 68.","Eigenmann, C. H. (1913) Some results from an ichthyological reconnaissance of Colombia, South America [Part II]. Indiana University Studies, (18), 1 - 32.","Ibarra, M. & Stewart, D. J. (1987) Catalogue of type specimens of recent fishes in Field Museum of Natural History. Fieldiana, Zoology, 35, 1 - 112.","Kramer, D. L. (1978) Reproductive seasonality in the fishes of a tropical stream. Ecology, 59 (5), 976 - 985. https: // doi. org / 10.2307 / 1938549","Kramer, D. L. & Bryant, M. J. (1995) Intestine length in the fishes of a tropical stream: 1. Ontogenetic allometry. Environmental Biology of Fishes, 42 (2), 115 - 127.","Bussing, W. A. (1987) Peces de las aguas continentales de Costa Rica. Editorial de la Universidad de Costa Rica ciudad Universitaria Rodrigo Facio, San Jose.","Bussing, W. A. (1998) Peces de las aguas continentales de Costa Rica. Editorial de la Universidad de Costa Rica ciudad Universitaria Rodrigo Facio, San Jose.","Dahl, G. (1971) Los peces del norte de Colombia. Bogota: Inderena."]}

Hyphessobrycon savagei Bussing, 1966 (Figs. 19���20, Tab. 6) Hyphessobrycon savagei Bussing, 1966: 215 [original description, type locality: ��� Puntarenas: R��o Ceibo (260m) at IH, 6 km W of Buenos Aires turnoff���]���. G��ry, 1977: 463 [diagnosis in key; within group ���b���]���. Bussing, 1987: 87 [Costa Rica; diagnosis in key, brief description, photo, distribution]���. Bussing, 1998: 108���111 [Costa Rica; diagnosis in key; brief description, photo, distribution]���.Lima et al., 2003: 140 [listed; distribution]���. Zarske & G��ry, 2002: 25 [diagnosis from Hyphessobrycon columbianus; key to species; color pattern in life]���. Angulo et al., 2013: 992 [listed, Costa Rica]���. Angulo et al., 2015: 376 [type catalog]. Diagnosis. Hyphessobrycon savagei differs from all congeners, except from H. compressus, H. diancistrus, H. otrynus, and those belonging to the H. panamensis species-group, by the presence of one large bony hook per fin ray, anterodorsally oriented and surrounded by a fleshy area, on last unbranched and first branched anal-fin rays of mature males (vs. absence of hooks, or presence of small and numerous hooks per fin ray). Hyphessobrycon savagei can be distinguished from H. compressus by the absence of a black blotch on the dorsal fin (vs. presence), by the presence of predorsal series (vs. absence), and by presenting 31���34 scales in longitudinal series (vs. 41���48). It can be distinguished from H. diancistrus and H. otrynus by presenting the caudal fin hyaline (vs. dark-brown blotches on caudal-fin lobes in H. diancistrus; distal third of caudal fin darkened in H. otrynus). It differs from the remaining species of the H. panamensis species-group by presenting two humeral blotches equally pigmented, extending two to four longitudinal scale rows below lateral line, with contours and margins always well-defined (vs. one diffuse humeral blotch, in H. daguae; second humeral blotch less conspicuous than the first, extending up to two longitudinal scale rows below lateral line, with posterior margin as a concentration of melanophores concentrated along longitudinal midlateral stripe, in H. bussingi, H. columbianus, H. condotensis, and H. panamensis). It can be further distinguished from H. columbianus by the absence of a silvery-blue iridescent coloration on dorsal region of flanks in life (vs. presence), and by a lower body depth (33.3���45.5% vs. 46.0���50.3% SL). It also differs from H. bussingi and H. condotensis by presenting 19���24 branched anal-fin rays (mode = 23) (vs. 23-27, modes = 24 and 25, respectively). It can be distinguished additionally from H. panamensis by presenting the first humeral blotch very conspicuous (vs. inconspicuous or less conspicuous). Description. Morphometric data in Table 6. Body compressed, greatest body depth slightly anterior to vertical through dorsal-fin origin. Dorsal profile of body slightly convex from upper lip to vertical through anterior nostril; straight or slightly convex from this point to distal tip of supraoccipital process. Slightly convex from tip of supraoccipital process to dorsal-fin origin; slightly convex from this point to adipose-fin origin; and slightly concave along caudal peduncle. Ventral profile of body convex from lower lip to anal-fin origin; roughly straight along anal-fin base; and slightly concave along caudal peduncle. Jaws equal, mouth terminal. Premaxillary teeth in two rows; outer row with 1(1), 2(6), 3(31), or 4(17) tricuspid teeth; inner row with 4(1), 5(42), 6(16), 7(1), or 8(1) conical to pentacuspid teeth. Posterior margin of maxilla extending to vertical through anterior margin of pupil. Maxilla with 2(4), 3(42), 4(13) conical, tri- to pentacuspid teeth. Dentary with 4(44), 5(13) large tri- to pentacuspid teeth, gradually decreasing in size, followed by 10 conical and abruptly smaller teeth. Central cusp longer than lateral ones. Scales cycloid, with few radii (4-10); circuli markedly anterior and marginally (dorsally and ventrally). Lateral line incompletely pored with 8(2), 9(6), 10(12), 11(21), 12(5), 13(7), 14(3), or 16(1) perforate scales. Longitudinal series, including perforate scales, with 31(13), 32(14), 33(25), or 34(3). Scale rows between dorsal-fin origin and lateral line 6(24), 6��(3), or 7(32). Scale rows between lateral line and pelvic-fin insertion 5(32), 5��(18), 6(26), or 6��(6). Predorsal series with 9(13), 10(33), 11(11), or 12(1). Single row of up to 13 scales covering base of anteriormost anal-fin rays. Circumpeduncular scales 13(5) or 14(42). Dorsal-fin rays ii(60) or iii(1), 8,i(7), or 9(53). First dorsal-fin pterygiophore located between neural spines of ninth and 10 th (2) vertebrae. Adipose fin present. Pectoral-fin rays i,11(6), 12(41), or 13(13), distal tip of rays reaching pelvic-fin origin. Pelvic-fin rays i,6(1), or 7(58), distal tip surpassing anal-fin origin. Anal-fin rays iii(4), iv(50), or v(6), 19(3), 20(9), 21(14), 22(9), 23(16), or 24(7). Anal fin of mature males with large bony hooks. Caudal fin bifurcate, lobes approximately of same size, i,9/8,i rays. First gill arch with 6(34), 7(21), 8(1) rakers on epibranchial, 1(60) on intermediate cartilage, 10(21), 11(34), 12(3), or 13(1) on cerato+hipobranchial. Branchiostegal rays 4. Total vertebrae 32(1), 33(1), supraneurals 4(1) or 5(1) (Fig. 20). Color in alcohol. Background coloration beige to yellowish. Dorsal portion of head and middorsal region of body with dense concentration of melanophores. Region above and below pupil darkened. Infraorbitals and opercle beige, with few scattered melanophores, evenly distributed. Lower portion of maxilla and gular region with few scattered melanophores, slightly more concentrated on upper half. Two dark-brown humeral blotches, vertically elongated, separated by light-beige area. First humeral blotch conspicuous, with melanophores slightly more concentrated on superior area, extending four to five longitudinal scale rows above and three to four below lateral line. Second humeral blotch conspicous, with melanophores equally distributed, extending three to four longitudinal scale rows above and one or two below lateral line. Body sides with melanophores evenly distributed, except on anteroventral half, with few or without melanophores. Brownish to silvery longitudinal midlateral stripe, from second humeral blotch to caudal peduncle, more conspicuous on posterior region. Fins hyaline, with few melanophores; anterior half of dorsal fin, distal third of anal fin, and median caudal-fin rays with more concentration of melanophores. Abdominal region, from pectoral-fin origin to pelvic-fin end, without melanophores. Color in life. Background color silvery. Dark markings as in preserved specimens. Region above anal fin purple iridescent. Dorsal and caudal fins orange. Pelvic and anal fins (especially next to hooks) bright red. Adipose fin yellow. Females with less intense color pattern (Bussing, 1966, 1998). Distribution. Hyphessobrycon savagei is known from the r��o P��rris and r��o T��rraba, Pac��fic coast of Costa Rica (Figs. 4 and 21). Sexual dimorphism. Mature males with one large bony hook by ray, antero-dorsally oriented and surrounded by fleshy area, from the last unbranched (on fifth or sixth segments) to the third branched (on the six segment) analfin ray of mature males (Figs. 5c and 6c). Small hooks on distal segments of anteriormost anal-fin rays were not observed. Mature males with distal margin of anal fin almost straight, and no evident elongation on dorsal fin, and females with distal margin of anal fin falcate, with anteriormost anal-fin rays longer than posterior ones, forming a lobe. Ecology and habitat. Hyphessobrycon savagei lives in coastal areas of streams and rivers in altitudes between 0-70 meters a.s.l., in both stagnant waters and rapids, with water temperatures ranging between 24-30��C. It feeds on aquatic and terrestrial insects (Bussing, 1998). Material examined. 180 specimens (23.8���42.1 mm SL). All from Costa Rica. Type material. UCR 42-1, paratypes of Hyphessobrycon savagei, 43 (rd, 15, 23.8 ��� 40.7 mm SL): San Jos��, creek Paja de Agua, 8 km Southern San Isidro del General, north side of road Interamericana, 09��19���10������N 83��39���54.1������W; W. A. Bussing & M. I. Bussing, 29 Jul 1966. Non-type material. Puntarenas, r��o T��rraba basin: ANSP 164253, 100 (30, 27.5-32.6 mm SL, 2 c&s, 29.2-29.3 mm SL), culvert pool at Inter American Highway, ca. 20 km S of Palmar Norte; D. Fromm, 10 Mar 1989; UCR 112012, 575 (40, 28.9 ��� 34.9 mm SL), creek 200 m Southeast of r��o Salam�� Nuevo on road Interamericana, 08��50���10������N 83��17���49.2������W; W.A. Bussing & O. Blanco, 05 Jan 1967; UCR 114014, 60 (25, 27.1 ��� 42.1 mm SL), creek 36, 12 km West of village r��o Claro on road Interamericana, 08��42���50������N 83��09���9.8������W; W.A. Bussing & O. Blanco 05 Jan 1967; San Jos��: UCR 40001, 54 (20, 25.9 ��� 29.7 mm SL), r��o Pedregoso, 3 km West of San Isidro del General on farm Villa Nueva, 09��22���35������N 83��43���44.2������W; O. Blanco, 19 Jul 1966; UCR 1942001, 139 (40, 25.8 ��� 32.5 mm SL), Corcovado National Park, r��o Camoral, next to Sirena station, 08��28���50������N 83��35���20������W; J. Lyons, 07 Mar 1987. Puntarenas, r��o P��rris basin: UCR 304002, 164 (40, 28.3 ��� 35.4 mm SL), tributary of the r��o Jicote, 0.5 km Southern of Loma, on road Puriscal-Parrita, 09��32���25������N 84��23���14.7������W; W. A. Bussing, R. Nishimoto, J. Perry & C. Mata, 16 Jan 1969., Published as part of Ota, Renata R., Carvalho, Fernando R. & Pavanelli, Carla S., 2020, Taxonomic review of the Hyphessobrycon panamensis species-group (Characiformes: Characidae), pp. 401-436 in Zootaxa 4751 (3) on pages 427-430, DOI: 10.11646/zootaxa.4751.3.1, http://zenodo.org/record/3714610, {"references":["Bussing, W. A. (1966) New species and new records of Costa Rican freshwaters fishes with a tentative list of species. Revista de Biologia Tropical, 14 (2), 205 - 249.","Gery, J. (1977) Characoids of the world. TFH Publications, Neptune City, 672 pp.","Bussing, W. A. (1987) Peces de las aguas continentales de Costa Rica. Editorial de la Universidad de Costa Rica ciudad Universitaria Rodrigo Facio, San Jose.","Bussing, W. A. (1998) Peces de las aguas continentales de Costa Rica. Editorial de la Universidad de Costa Rica ciudad Universitaria Rodrigo Facio, San Jose.","Zarske, A. & Gery, J. (2002) Der Blaurote Kolumbien-Salmler. Hyphessobrycon columbianus n. sp. - ein neuer Salmler (Teleostei; Characiformes, Characidae) aus dem kolumbianischen Darien. Das Aquarium, 36 (391), 22 - 30.","Angulo, A., Garita-Alvarado, C., Bussing, W. A. & Lopez, M. I. (2013) Annotated checklist of the freshwater fishes of continental and insular Costa Rica: additions and nomenclatural revisions. Check List, 9 (5), 987 - 1019. http: // doi. org / 10.15560 / 9.5.987","Angulo, A., Arias-Godinez, G., Lopez, M. & Bussing, W. (2015) Catalogo de material tipo depositado en la coleccion ictiologica del museo de zoologia de la universadad de Costa Rica. In: Del Moral Flores, L. F., Ramirez Villalobos, A. J., Martinez Perez, J. A., Gonzalez Acosta, A. F. & Franco Lopez, J. (Eds), Colecciones Ictiologicas de Latinoamerica. Universitad Nacional Autonoma de Mexico, Ciudad de Mexico, pp. 365 - 386."]}

Hyphessobrycon itaparicensis Lima & Costa, 2001 Figs. 1-4, Tab. 1 Hyphessobrycon itaparicensis Lima & Costa, 2001: 233-234; 236-237. Type locality: small stream in Ilha de Itaparica. - Lima et al., 2003: 139 (list of species). - Buckup et al., 2007: 54 (list of species). - Menezes et al., 2007: 90 (list of species). - Zanata, Camelier, 2010: 771-772; 776 (diagnosis of H. brumado; comparative material). - García-Alzate et al., 2010: 58; 62 (diagnosis and comparison with H. sebastiani) - Burger et al., 2011: 274; 277; 279; 282; 285; 290 (occurrence of species on the Recôncavo Sul basin, Bahia State; photo; identification key; comparative material). - Carvalho, Langeani, 2013: 533 (comparative material). - Menezes et al., 2013: 29 (comments about conservation of type locality). - Brito et al., 2014: 1156-1159 (geographic distribution; photos; molecular identification). - Camelier, Zanata, 2014: 687; 690; 691; 692; 696 (list of species; geographic distribution; comments). - Dagosta et al., 2014: 373 (comparative material). - Carvalho et al., 2014: 248 (diagnosis of H. flammeus). - Lima et al., 2014: 170 (diagnosis of H. montagi). - Vieira et al., 2016: 57- 60 (description of pelvic- and anal-fins bony hooks; examined material). Hyphessobrycon cf. itaparicensis. - Costa, 2004: 6 (habitat notes). Hyphessobrycon ellisae Bragança, Ottoni & Rangel-Pereira, 2015: 256 (original description, type locality: Brazil, Sergipe State, Município de Estância, about 8 km north of Santa Cruz do Abais). Preoccupied by H. ellisae (Pearson, 1924) and replaced by Hyphessobrycon sergipanus (Bragança et al., 2016). Hyphessobrycon sergipanus Bragança, Ottoni & Rangel- Pereira, 2016: 373 (name replacement for H. ellisae). NEW SYNONYM. Diagnosis. Hyphessobrycon itaparicensis can be easily distinguished from most congeners, except the species of the “rosy tetra clade” sensu Weitzman, Palmer (1997), and H. balbus Myers, H. bifasciatus Ellis, H. chocoensis García- Alzate, Román-Valencia & Taphorn, H. columbianus Zarske & Géry, H. eilyos Lima & Moreira, H. flammeus Myers, H. gracilior Géry, H. griemi Hoedeman, H. igneus Miquelarena, Menni, López & Casciotta, H. panamensis Durbin, H. savagei Bussing, H. scutulatus Lucena, H. sebastiani García-Alzate, Román-Valencia & Taphorn, H. taguae García-Alzate, Román-Valencia & Taphorn, H. tortuguerae Böhlke, and H. weitzmanorum Lima & Moreira, by the absence of any concentration of dark chromatophores at the caudal-peduncle region (vs. presence of a dark blotch or a longitudinal dark stripe extending over the caudal-peduncle region). Among the “rosy tetra clade”, H. itaparicensis differs from H. axelrodi (Travassos), H. bentosi Durbin, H. compressus (Meek), H. copelandi Durbin, H. dorsalis Zarske, H. epicharis Weitzman & Palmer, H. eques (Steindachner), H. erythrostigma (Fowler), H. georgettae Géry, H. haraldschultzi Travassos, H. heteresthes Ulrey, H. jackrobertsi Zarske, H. khardinae Zarske, H. megalopterus (Eigenmann), H. micropterus (Eigenmann), H. minor Durbin, H. pando Hein, H. pyrrhonotus Burgess, H. rosaceus Durbin, H. roseus (Géry), H. simulatus (Géry), H. socolofi Weitzman, H. sweglesi (Géry), and H. takasei Géry by the absence of a black dorsalfin blotch (vs. presence). The species can be diagnosed from H. bifasciatus, H. flammeus, H. griemi, H. savagei, H. sebastiani, H. tortuguerae, and H. weitzmanorum by the absence of a conspicuous second humeral blotch, having instead an inconspicuous concentration of melanophores without defined limits, that initiates immediately posterior to a clear area on the rear of the first humeral blotch and fades posteriorly (vs. second humeral blotch as conspicuous as first humeral blotch). The species can be further distinguished from some aforementioned species by presence of 3-11 maxillary teeth (vs. 1-2 in H. bifasciatus, H. balbus, H. igneus, and H. ecuadoriensis, and 2 in H. chocoensis and H. sebastiani), 5-8 pored scales on lateral line (vs. 9-13 in H. columbianus, and 10-13 in H. savagei), presence of humeral blotch (vs. absence in H. eilyos, H. gracilior, and H. scutulatus), caudal fin hyaline (vs. caudal fin with black median stripe in H. weitzmanorum), 32-36 longitudinal scales series and vertically elongated humeral blotch (vs. 29-31 longitudinal scales series and rounded humeral blotch in H. taguae), and maxillary teeth tricuspid (vs. pentacuspid teeth in H. panamensis). When alive, H. itaparicensis can be diagnosed from most congeners by usually having yellowish body and fins, allied to a dark brown or reddish midlateral stripe extending from dorsalfin base to caudal peduncle. Description. Morphometric data are summarized in Tab. 1. Body somewhat compressed and elongate. Greatest body depth at vertical through dorsal-fin origin or slightly ahead of this point. Dorsal profile of head somewhat convex from upper lip to vertical through anterior nostrils; straight to slightly convex above eye and somewhat concave from the vertical through posterior border of eye and to tip of supraoccipital spine. Dorsal profile of body somewhat convex from tip of occipital spine to dorsal-fin origin; straight to somewhat convex and posteroventrally slanted along dorsal-fin base; straight from end of dorsal-fin base to adipose fin and slightly concave along caudal peduncle. Head rounded anteriorly in lateral profile. Ventral profile of head and body convex from lower lip to pelvic-fin origin; straight from that point to anal-fin origin; straight and posterodorsally slanted along anal-fin base, and slightly concave along ventral profile of caudal peduncle. Eyes relatively large compared with head length. Lower jaw slightly longer than upper jaw, mouth terminal. Posterior terminus of maxilla usually extending beyond vertical through anterior margin of orbit. Nostrils close to each other and separated by skin flap; anterior opening small, semicircular and with dermal flap; posterior one more than twice in size, elongate and without dermal flap. Nasal bone present. Frontals separated anteriorly, with wide fontanel; parietal fontanel large, extending from epiphyseal bar to supraoccipital spine, slightly narrowed anteriorly. Infraorbital series variable; usually six elements but only five elements are present in some specimens, possibly due to fusion of infraorbitals three with four (e.g., UFBA 7553, 27.4 mm SL). Third infraorbital largest and contacting laterosensory canal of preopercle ventrolaterally. Laterosensorial canal of first infraorbital absent; canal present and close to inner margin of orbital rim from second to fourth infraorbitals. Degree of development of the parietal branch of the supraorbital canal variable, reaching parietal bone (UFBA 7558), just reaching suture between frontal and parietal bones (CIUFS 426, 38.9 mm SL; UFBA 7553, 27.4 mm SL), or poorly developed, not reaching the suture between frontal and parietal bones (CIUFS 694, 21.9 mm SL; UFBA 7515, 19.0 mm SL). Premaxillary teeth in two rows; outer row with one (2), two* (46) or three (52) teeth bearing three cusps; inner row with four (1), five* (74), six (24), or seven (1) teeth bearing three to five cusps; symphyseal tooth of inner series narrow, asymmetrical, usually without cusp on anteromedial side, one larger central cusp and one smaller on lateral side; second or third tooth the largest, with three or five cusps; last teeth smaller with three cusps or conical. Maxilla with three* (11), four (25), five (16), six (26), seven (12), eight (2), nine (4), or 11(1) teeth; anterior ones usually with three cups and posterior ones conical. Dentary with 11(1), 12(5), 14(2), 15(2), 16(1), or 19(1) teeth; four or five large anterior teeth cusps with three in most specimens examined, but specimens around 37.0 mm SL or larger with anterior teeth pentacuspid; seven to 14 smaller posterior teeth, usually conical (Fig. 2). Scales cycloid, circuli absent on exposed area of scales, with several parallel radii extending to posterior margin of scale. Lateral line incomplete; with five (3), six* (38), seven (42), or eight (10) pored scales; longitudinal scales series including pored scales 32(1), 33(10), 34(22), 35(19), or 36(3). Some scales lost in the holotype. Horizontal scale rows between dorsal-fin origin and pelvic-fin insertion 12*(14) or 13(78), commonly six above and six below lateral line, and more rarely seven above and six below. Scales along middorsal line between tip of supraoccipital process and origin of dorsal fin 10*(36), 11(35), or 12(9). Horizontal scale rows around caudal peduncle 13*(5) or 14(53). Base of anteriormost anal-fin rays covered by a series of three to five scales. Caudal fin with scales restricted to the base of rays. Muscular reduction at vertical through anterior portion of the swimbladder; swimbladder not completely exposed but covered by a thin layer of musculature; muscular reduction between first and second pleural ribs, visible or not by transparency through the body wall as a deep dark area under the humeral blotch. Dorsal-fin rays ii,8(3), ii,8,i(6), ii,9*(82), iii,9(4), ii,10(7). Distal margin of dorsal fin straight or slightly rounded. Dorsal-fin origin situated at vertical through approximately middle of standard length. Base of last dorsal-fin ray anterior to vertical through anal-fin origin. First dorsal-fin pterygiophore inserting behind neural spine of 10 th (8) vertebra. Adipose fin present. Anal-fin rays unbranched rays iii*(24), iv(70) or v(1); branched rays 20(2), 21(19), 22(24), 23*(28), 24(17), 25(7), or 26(1). Distal margin of anal fin slightly concave. First anal-fin pterygiophore inserting behind haemal spine of 15 th (4) or 16 th (4) vertebra. Pectoral-fin rays i,9(11), i,9,ii(3), i,10(46), i,10,i(1), i,11*(36), or i,12(5). Tip of pectoral fin usually reaching or slightly surpassing vertical through pelvic-fin insertion. Pelvic-fin rays i,5(1), i,6*(102), or i,7(2); tip of pelvic fin of mature males usually extends beyond insertion of first anal-fin ray. Caudal fin forked, lobes pointed, similar in size. Principal caudal-fin rays i,9+8,i(11). Eight (4), nine (4), 10(1), or 11(1) dorsal procurrent caudal-fin rays, and eight (7) or nine (3) ventral procurrent caudal-fin rays. First gill arch with five (4), six (3), or seven (1) gill rakers on epibranchial, eight (2) or nine (6) on hypobranchial and ceratobranchial, and one (8) on cartilage between ceratobranchial and epibranchial. Precaudal vertebrae 13(2), 14(7), or 15(2) and caudal vertebrae 19(1), 20(6), 21(3), or 22(1); total vertebrae 33(1), 34(8), 35(1), or 37(1). Supraneurals four (2), five (6), or six (2). Branchiostegal rays four (11). Coloration in alcohol. Overall ground color yellow to light brown (Fig. 1). Guanine restricted to part of infraorbitals, preopercle, and opercle in somewhat recently fixed specimens. Dorsal part of head with melanophores sparsely and evenly distributed, usually darker posterior to eyes. Melanophores sparsely distributed over maxilla and lateral portion of head, larger posterior to eyes. Opercle with sparse melanophores, usually more concentrated on its dorsal half. Ventral portion of head with a few scattered small melanophores, more concentrated on anteriormost portion. Dorsum and scales along lateral of body with sparse melanophores; middorsal series usually homogeneously darkened; two or three dorsalmost series of scales with clearer posterior border. Scales below it with melanophores homogeneously distributed. Abdominal region clear or with sparse tiny melanophores. Humeral region with a vertically-elongated faint humeral blotch, wider dorsally and tapering ventrally; widest portion located on second horizontal series above lateral line, reaching three or four scales horizontally. Humeral blotch preceded and followed by clear areas, although clear areas not completely devoid of melanophores; blotch formed by superficial melanophores. Some specimens, usually about 33.0 mm SL or larger, with concentration of melanophores posterior to the clear area on the rear of the humeral blotch, but not characterizing a well-defined second blotch (see item Discussion). Midlateral black narrow stripe along horizontal septum from vertical through dorsal-fin origin to caudal peduncle, more evident in the stretch posterior to the dorsal-fin origin, but falling short of the end of caudal peduncle; dark line or stripe formed by melanophores over skin and also embedded dark pigmentation; largest specimens (about 37.0 mm SL or larger) with longitudinal stripe slightly wider and less conspicuous. No distinct caudal-peduncle blotch or/and stripe. All fins slightly darkened; dorsal, caudal, and pectoral usually with tiny melanophores forming dark lines along borders of rays and sparse melanophores on interradial membranes. Anal fin similarly colored, but lacking dark lines on borders of rays; distal border of fin somewhat darker, more evident on anteriormost rays. Pelvic fin somewhat less colored, with melanophores usually restricted to distal portion of rays; some specimens with distal half of rays distinctly dark. Adipose fin mostly hyaline; some specimens with few scattered small melanophores at base of fin. Specimens sampled in black water streams usually with overall darker coloration of body and fins, rendering some patterns and blotches described above somewhat merged with ground coloration and inconspicuous. Coloration in life. Life color pattern is somewhat variable along distribution and apparently influenced by the physical characteristics of water body inhabited by each population (Fig. 3). Specimens from clear water rivers usually possess yellowish overall body coloration, including distinct yellow coloration of fins, maxillae, and dorsal half of head (Figs. 3a, c-e). Additionally, a brown-reddish longitudinal stripe, from the rear of the humeral blotch or body midlength to the caudal peduncle is observed in some of those specimens (Fig. 3d). On the other hand, specimens from dark waters are more pigmented, with no humeral blotch or clear surrounding areas visible (Fig. 3b). These specimens are usually shiny silver, with large amount of guanine over scales on flank and dark longitudinal stripe, when visible. They possess yellow or orange chromatophores over scales on anterior half of body and strong yellow to orange fins. Sexual dimorphism. Lima, Costa (2001: 235) mentioned “no hooks on fins” of H. itaparicensis. However, Brito et al. (2014) and Vieira et al. (2016) recently reported the presence of bony processes on anal and pelvic fins in H. itaparicensis specimens from rivers draining Sergipe State. Examination of paratypes of H. itaparicensis (MZUSP 57540, UFRJ 4843) and various recently sampled specimens also revealed well-developed bony processes on first to fourth branched anal-fin rays and on the anteriormost two or three branched pelvic-fin rays (Fig. 4). Anal-fin bifurcated bony processes are distributed from the last unbranched up to the eighth branched anal-fin rays, usually on distal half of rays, on the segment just before the bifurcation of rays and continuing on dorsal hemitrichium of each ray. Bony processes are concave, anterodorsally directed bilaterally, symmetric, larger around midlength of branched portion of rays (Fig. 4a), and usually more numerous in the third and fourth branched anal-fin rays, with up to 12 paired processes (UFBA 7558, 40.5 mm SL). Pelvic-fin rays have similar concave and anterodorsally directed bony processes on the first and second, rarely on third, branched rays of mature males (Fig. 4b; see also Brito et al. (2014: 1159, Fig. 5d). However, pelvic-fin bony processes are not bifurcate and are distributed on the border of three or four segments anterior to branching point or restricted to the dorsal hemitrichium. Bony processes decrease in size on distal portion of rays. Up to 15 bony processes were observed in the first and 13 processes on the second branched ray (UFBA 7558, 40.5 mm SL). In one paratype (MZUSP 57540, 25.4 mm SL), six processes were observed in each of the two first branched rays and in another paratype (UFRJ 4843, 32.0 mm SL), processes occur on three first branched rays. Other sexually dimorphic traits observed in H. itaparicensis include interradial membrane on areas with bony processes on both fins tumescent, shape of the analfin profile distinct in males and females, and pelvic-fin length distinct in males and females. According to Brito et al. (2014: 1159, Figs. 5 a-b) the anal-fin distal profile is almost straight in males vs. concave from fifth to tenth branched ray in females. Examination of mature males performed herein corroborates this information, although with some variation among populations examined. Brito et al. (2014) described the distal end of the pelvic fin surpassing the anal-fin origin in males but not reaching the anal fin in females. However, examination of paratypes (UFRJ 4843, 32.0) revealed pelvic fin of females reaching the anal-fin origin and non-type mature males with pelvic fin barely reaching anal-fin base (UFBA 7558, 40.5 mm SL). Males usually have tips of pelvic-fin rays reaching beyond the anal-fin origin and overlapping the base of the first branched fin rays, while in females the pelvic fin may reach the anal fin but does not reach the basal portion of the branched fin rays. The holotype is apparently a female, relatively large-sized, without bony processes on fin rays, with concave anal-fin profile, and pectoral-fin barely reaching pelvic-fin insertion. Distinct elongation of fins in males and sexually dimorphic coloration typical of various species of Hyphessobrycon were not observed in H. itaparicensis. Gill glands (Burns, Weitzman, 1996) were not found on first gill arch of neither sex. Geographic distribution. Hyphessobrycon itaparicensis occurs in small coastal Brazilian rivers in the Northeastern Mata Atlântica freshwater ecoregion (NMAF), from streams around Camamu municipality, Bahia State in its southernmost distribution, to tributaries of rio Sergipe in Areia Branca municipality, Sergipe State, up north (Fig. 5). The known distribution of this species is restricted to a group of basins proposed by Camelier, Zanata (2014), the ‘Group North’, which includes drainages situated in the northernmost portion of the NMAF ecoregion. Ecological notes. Hyphessobrycon itaparicensis was first sampled in a small clear water stream (about 4 m wide and 1 m deep) below a small waterfall in Ilha de Itaparica, located about 200 m from the sea, although without tidal influence (Lima, Costa, 2001), in Barra do Gil, municipality of Vera Cruz (C. Sampaio, person. comun.), on the eastern portion of the island. According to

Hyphessobrycon flammeus Myers, 1924 Figs. 1-5 Hyphessobrycon flammeus Myers, 1924: 330-331. Type locality: Rio de Janeiro, Brazil. - G��ry, 1977: 458 (classification in artificial group b); 463 (key for the species); 465 (image). -Weitzman & Vari, 1988: 447; 450 (listed as miniature species). -Weitzman et al., 1988: 419-420 (notes about geographic distribution and biogeography). - Mazzoni et al., 2000: 66 (citation for areas of occurrence - fluvial lowland and marsh). -Lima et al., 2003: 137 (check list for the genus). -Lima & Moreira, 2008: 72-73 (threatened species in Brazil, category EN - A2ace; B2ab(iii); historic and biology notes; additional notes about geographic distribution, biogeography and conservation strategies). -Oyakawa et al., 2009: 366 (listed as threatened for S��o Paulo State, category EN B2abiii; comments about distribution, ecological notes). - Marceniuk et al., 2011: 221; 223-225; 232; 235 (listed). Diagnosis. Hyphessobrycon flammeus differs from its congeners (except H. bifasciatus, H. balbus Myers, H. chocoensis Garc��a-Alzate, Rom��n-Valencia & Taphorn, H. columbianus Zarske & G��ry, H. condotensis Regan, H. griemi Hoedeman, H. igneus Miquelarena, Menni, L��pez & Casciotta, H. itaparicensis Lima & Costa, H. panamensis Durbin, H. savagei Bussing, H. sebastiani Garc��a-Alzate, Rom��n-Valencia & Taphorn, H. tortuguerae B��hlke, and H. weitzmanorum Lima & Moreira) by presenting two humeral spots vertically elongated and no caudal peduncle blotch. Hyphessobrycon flammeus differs from aforementioned species by presence of 5-8 maxillary teeth (vs. 1-3 in H. balbus, 1-2 in H. bifasciatus and H. igneus, 3 in H. condotensis and H. panamensis, 2-3 in H. griemi, 1-4 in H. savagei, 2 in H. chocoensis and H. sebastiani, 9-10 in H. tortuguerae), caudal fin hyaline (vs. caudal fin with black median stripe in H. weitzmanorum), no longitudinal stripe dark and second humeral spot conspicuous as well as first humeral spot (vs. faint longitudinal stripe dark and second humeral spot less defined than first in H. weitzmanorum). Description. Morphometric data are summarized in Table 2. Body compressed, moderately short, greatest body depth at vertical through dorsal-fin origin. Dorsal profile of head convex from tip of upper jaw to vertical through anterior nostril; slightly straight or concave from that point to tip of supraoccipital spine. Dorsal profile of body slightly convex from posterior tip of supraoccipital spine to base of last dorsal-fin ray, and straight to adipose-fin origin; last dorsalfin ray at vertical through of anal-fin base. Ventral profile of body convex from tip of lower jaw to pelvic-fin origin, straight or slightly convex from that point to anal-fin origin, and straight and/or subtly rounded along anal-fin base. Dorsal and ventral profiles of caudal peduncle slightly straight. Eyes relatively large compared with head, without distinct adipose eyelid. Lower jaw slightly longer than upper jaw, mouth slightly subterminal. Maxilla extending posteriorly surpassing anterior margin of orbit to vertical through crystalline lens anterior margin, slightly curved, aligned approximately at 45 degrees angle relative to longitudinal axis of body. Nostrils close to each other, anterior opening small and circular, posterior one twice in size and slightly reniform or elongate. Nostrils separated by skin flap. Nasal bone absent. Frontals separated anteriorly, with wide fontanel; parietal fontanel large, extending from epiphyseal bar to supraoccipital spine, slightly narrowed anteriorly. Infraorbital series with four or five elements (probably fifth and sixth, when present, fusioned). Laterosensorial canal of first infraorbital absent, from second to fourth canal close to inner margin of orbital rim. Third infraorbital largest, double size compared to other (in length and depth) and contacting laterosensory canal of preopercle ventrolaterally. Premaxillary teeth in two rows: outer row with 1*(12) or 2(46) tricuspid teeth; inner row with 5*(57) or 6(1) pentacuspid teeth. Dentary with 4*(59) or 5(1) large, tri- to pentacuspid teeth, followed by 4(1), 5(14), 6(38), 7(6), or 8(1) small, conical or tricuspid, teeth abruptly smaller than anterior largest teeth. Dorsal border of maxilla relatively straight. Maxilla with 5*(35), 6(13), 7(10), 8(4), or 9(1) conical, tri- to pentacuspid teeth along anteroventral margin (Fig. 3). Central median cusp of all teeth longer than remaining lateral cusps; cusp tips slightly curved inward on dentary teeth, and relatively straight on premaxillary teeth. Scales cycloid, with few and relatively small radii (3- 7); circuli marked anteriorly and marginally (dorsal and ventral). Lateral line incomplete, extending to vertical through middle of pectoral fin; perforated scales 4*(23), 5*(24), or 6(9); longitudinal scales series including pored scales 26(8), 27(1), 28(17), 29(6), 30*(15), 31(4), 32(1), 33*(4), or 34(2); five*(52) or six*(6) scale rows between dorsal-fin origin and lateral line; five*(3) or six*(55) scale rows between lateral line and pelvic-fin origin. Predorsal scales 6(1), 7(1), 8*(27), 9*(14), or 10(15). Scale sheath along anal-fin base in single row of 4(13), 5*(25), 6(11), or 7(6) scales. Circumpeduncular scales 14*(47) or 15(6). Axillary scale absent. Dorsal-fin rays ii,9*(62) or 10(3); first unbranched ray approximately one-half of second one or shorter.Dorsal-fin origin at midbody or posterior, at vertical through anterior third of pelvic fin base. Base of last dorsal-fin ray at vertical through distal tip of pelvic fin, anterior to origin of anal fin. Tip of longest ray of adpressed dorsal fin at vertical through base of first two branched anal-fin rays. First dorsal-fin pterygiophore inserting between neural spine of 10 th (1) and 11 th (3) precaudal vertebra. Adipose fin present. Pectoral fin with i,9*(2), 10*(40), or 11(11) rays, extending to pelvic fin origin. Pelvic fin with i,6*(43) or 7(22) rays; its origin anterior to vertical through dorsal-fin origin;tip of rays reaching anal-fin origin. Anal-fin rays iv,20(1), 21(3), 22*(11), 23(15), 24(21), or 25(8). Anal fin of males with small bony hooks. Caudal fin forked, lobes slightly rounded, similar in size, i,9/7,i (2), i,9/8,i*(41), or i,10/8,i (1) [one specimen with i,5/5,i, probably anomalous] rays. Caudal fin naked, scales restricted to its base. Dorsal procurrent caudal-fin rays 8(2) or 9(2) and ventral procurrent caudal-fin rays 7(1) or 8(3). Branchiostegal rays 4. First gill arch with 6(3) or 7(1) gill rakers on epibranchial, 1(4) between epibranchial and ceratobranchial, 8(3) or 9(1) on ceratobranchial, and 2(4) on hypobranchial. Precaudal vertebrae 13(2) or 14(2) and caudal vertebrae 17(1), 18(2), or 19(1); total vertebrae 31(1) or 32(3). Supraneurals 4(2) or 5(2), filiform, some with dorsal portion expanded. Color in alcohol. Overall body color yellowish to whitish. Dark chromatophores scattered on the lateral portion of head, more densely concentrated from snout to the supraoccipital. Second and third infraorbitals and upper region of opercular apparatus denser chromatophores, and yellowish adjacent areas. Small chromatophores scattered on the jaws. Two conspicuous vertically elongated humeral spots, extending from the dorsal portion to the pectoral fin, positioned over three to four vertical series of scales above the lateral line and four to five below it, and vertically extending over two to four horizontal series of scales, more intense in the middle of the humeral spot; both spots are separated by two to three horizontal scales. Abdominal region yellowish, with few scattered chromatophores. Lateral side of the body scattered melanophores, dorsal portion with a reticulated pattern on the posterior margin of its scales. Midlateral body stripe absents. All fins hyaline or slightly dark (see more details for males in sexual dimorphism). Color in life. Overall, body with an intense reddish color (hence the name flammeus, from latin, red - flame-colored). More intense concentration of erythrophores between verticals from the dorsal and/or pelvic-fin to anal-fin end. Distribution of melanophores on its head, flank and fins similar to the one described in alcohol coloration. Dorsal and dorsolateral regions of body rosaceous and orangish. Ventral area of its head up to its posterior humeral spot, including the abdominal region, slightly grayish to yellowish. Sometimes, the upper area of its head up to its first humeral spot with iridescent blue hues. All fins reddish, no pigmentation on the middle caudal-fin rays; caudal and anal fins with a more intense color. Males with black pelvic and anal fins tips and whitish dorsal fin (vs. absent in females) (Fig. 4). Sexual dimorphism. Males of Hyphessobrycon flammeus present bony hooks in the anal and pelvic fins (Fig. 5) (vs. absent in females). Small, simple bony hooks in the anal fin, extending from the last unbranched ray to the last branched rays, i.e., in all fin rays (Fig. 5a for the first bony hooks in the anal fin); pelvic fin with hooks in the first two or three branched rays (Fig. 5b). Distal portion of its anal fin slightly straight in males (vs. anteriorly falcate in females) (Fig. 2). Males normally smaller than females, but present a bright reddish color, with black tips on pelvic and anal fins. Darkish dorsal fin in the middle portion and whitish on the tip in both males and females. Gill glands (Burns & Weitzman, 1996) were not found macroscopically on first gill arch on both sexes. Distribution. Hyphessobrycon flammeus has occurrence in streams and coastal rivers of Rio de Janeiro State, in Guanabara bay basin, middle rio Para��ba do Sul basin and rio Guandu basin. In the upper rio Tiet�� drainage (upper rio Paran�� basin, S��o Paulo State) it is found around of the metropolitan region of S��o Paulo city (Fig. 6). Ecological notes. Hyphessobrycon flammeus has been reported in small streams of slow flowing, dark-brown or clear water, shaded by small forest (Myers, 1944). Recently, it has been found in upper rio Tiet�� drainage, in small streams of clear water and sandy bottom, as well as in the marginal portions of rio Tiet�� (Lima & Moreira, 2008). Life expectancy is around four years. In its natural habitat it forms relatively numerous schools with agonistic interaction among alpha males. The species prefer environments with vegetation and streams with slow flowing water, living in depths not superior to 50 cm, and water temperature from 22��C to 28��C. Its diet includes small insects, ���worms���, and plants (Miranda et al., 2012). The c&s stomach contents of four specimens contained aquatic insects, mainly Chironomidae larvae, and fine organic matter. During reproduction, the female deposits around 200 to 330 oocytes on rocks, plants, or submerged debris; after that, the male releases the sperm for fecundation. The eggs hatch in approximately 2-3 days (Miranda et al., 2012). Further ecological/biological information can be found in Myers (1924, 1945), Lima & Moreira (2008) and Oyakawa et al. (2009). On the Wide World Web, there is plenty of information shared among aquarists about behavior, reproduction, and many other biological characteristics of H. flammeus, including information on keeping and reproducing them in captivity. Conservation remarks. Hyphessobrycon flammeus has been in the Brazilian list of threatened fish species since 2004 (Di��rio Oficial da Uni��o, 2004), category EN-A2ace; B2ab(iii) (i.e., endangered species). However, after the last evaluation conducted by ICMBio (Instituto Chico Mendes de Conserva����o da Biodiversidade) (in 2012, unpublished data) the category EN will be maintained, but the criteria will be changed. Notwithstanding its relative abundance in upper rio Tiet�� drainage streams, it is not common in the region of Rio de Janeiro State as referred in the material which was examined herein and mentioned by Lima & Moreira (2008). Therefore, in order to preserve its original population, it will continue as a threatened species, and public policies should prioritize its maintenance and conservation. Popular names. Due to its popularity in the aquarium trade, Hyphessobrycon flammeus is known by many different names in the world, such as ���engra��adinho���, ���lambarizinho vermelho���, ���Rio tetra���, ���tetra-rio���, ���tetra-rosa���, ���tetra-vermelho���, and ���vermelhinho��� (Brazil), flame tetra (USA and Philippines), ���ognennaya tetra��� (Russian Federation), ���punatetra��� (Finland), ���red tetra��� (USA), ���r��d Rio��� (Denmark), ���roter von Rio��� (Germany) (Lima & Moreira, 2008; Froese & Pauly, 2013). Phylogenetic reconstruction. Phylogenetic analysis, according to implied weighting procedures (Mirande, 2009, 2010), with consensus of ���k��� = 9.99804 (three trees) and ���k��� = 10.97824 (three trees) values, resulted in one tree with 2209 steps, consistency index (CI) of 0.17 and retention index (RI) of 0.66. In this hypothesis, H. flammeus is the basal species of the clade 196 of Mirande (2010), i.e., (H. flammeus ((Aphyocharacinae (Aphyoditeinae, Cheirodontinae)) Stevardiinae)). Synapomorphies for this clade are: ch. 53:1 - length of supraoccipital spine extends only to anterior limit of neural complex; ch. 266:0 - two dorsal-fin rays articulating with first dorsal pterygiophore; ch. 363:1 - number of 2n chromosomes, 52 or more.Autapomorphies for H. flammeus are: ch. 33:1 - nasal bone absent; ch. 58:1 - bony lamellae bordering laterosensory canal of first infraorbital absent; ch. 99:0 - posterior extent of maxilla not reaching second infraorbital; ch. 168:2 - foramen in posterior region of metapterygoid in form of incomplete arch, bordered posteriorly by hyomandibula; ch. 201:1 - denticles on gill rakers absent; ch. 253:1 - position of ventral end of posttemporal posterior to lateral margin of epioccipital; ch. 305:0 - anterior ventral procurrent caudal-fin rays paired, only distally fused; ch. 332:1 - posterior attachment of A1 section of adductor mandibulae restricted or almost restricted to horizontal arm of preopercle; ch. 342:1 - second humeral spot present as a conspicuous vertical bar. Examined material. Syntypes. USNM 92969, 2 (2), 24.1-25.4 mm SL, Brazil, Rio de Janeiro State, vicinity of Rio de Janeiro, R. Brooca [probable], aquarium bred, no date. Non-types: All from Brazil. Rio de Janeiro State: Guanabara bay basin. MNRJ 8795, 16, 8.6-19.2 mm SL, floodplain near to Imbari��, Baixada Fluminense, L. Travassos, H. S. Lopes & H. Travassos, 04 Aug 1954. Middle rio Para��ba do Sul basin. MNRJ 19543, 20, 14.3-17.1 mm SL, 47 older road Rio-S��o Paulo, 15 Dec 1972. Rio Guandu basin: MZUSP 51020, 1(1), 17.8 mm SL, Itagua��, ribeir��o da Ponte do Teixeira, on road Itagua��/Raiz da Serra, Km 5, 08 Sep 1969. ZUEC 4253, 7, 15.7-20.4 mm SL, Itagua��, 15 Dec 1972. S��o Paulo State: rio Tiet�� drainage, upper rio Paran�� basin. LBP 8890, 1(1), 21.0 mm SL, Sales��polis, rio Tiet��, 10 Sep 2009. LBP 8905, 31 of 45 (31), 16.6-21.8 mm SL, Biritiba Mirim, rio Tiet��, 10 Sep 2009. MCP 20235, 1, 19.5 mm SL, Mogi das Cruzes, Tiet��, rio Taia��upeba near electric plant of Tijuco Preto in Taia��upeba, 12 Jan 1997. MCP 20237, 7 of 15, 13.2-20.9 mm SL, Biritiba Mirim, first stream in road from Biritiba Mirim to Casa Grande, 13 Jan 1997. MCP 20239, 3, 18.9-20.8 mm SL, Mogi das Cruzes, stream on road of Mogi das Cruzes to Sales��polis, ca. 8 km of Mogi das Cruzes, 13 Jan 1997. MCP 25545, 2, 15.5- 25.4 mm SL, Biritiba Mirim, Mogi das Cruzes, stream on road to Biritiba Mirim/Casa Grande, affluent of Ponte Nova Dam in the rio Tiet��, 17 Dec 1999. MNRJ 39230, 1, 26.9 mm SL, Mogi das Cruzes, district of Natureza de Taia��upeba, headwater of rio Grande, 07 Nov 2011. MNRJ 39235, 21, 14.2-29.4 mm SL, Mogi das Cruzes, stream tributary of rio Jundia�� together junction of road for fazenda rio Grande with municipal road Taia��upebaBertioga, 07 Nov 2011. MNRJ 39240, 31, 11.8-23.6 mm SL, Mogi das Cruzes, headwaters of rio Jundia��, fazenda do Sr. Celso Meida, 07 Nov 2011. MZUSP 86925, 6 of 7 (6), 16.2-18.2 mm SL, Biritiba Mirim, in the area of VCN Mining, 21 Mar 2005. MZUSP 88175, 1 (1), 12.6 mm SL, Embu, stream affluent of rio Embu with empties in reservoir of Guarapiranga, 23 Aug 2005. MZUSP 88183, 17, 14.0- 24.4 mm SL, Itapecerica da Serra, stream affluent of rio Embu-Mirim, near deposit of construction material Lagoa, on road Jo��o Rodrigues de Morais, 23 Aug 2005. MZUSP 88190, 1 (1), 13.5 mm SL, Itapecerica da Serra, stream affluent of rio Embu-Mirim, on road Jo��o Rodrigues de Morais, Lagoa neighborhood, 23 Aug 2005. MZUSP 88198, 1 (1), 17.1 mm SL, Itapecerica da Serra, stream affluent of rio Embu-Mirim, Lagoa neighborhood, 23 Aug 2005. MZUSP 88207, 19 (10, 1 c&s, 11.2- 19.6 mm SL), 9.6-19.6 mm SL, stream affluent of Guarapiranga dam, on road of Jaceguava, near soccer stadium, Casa Grande neighborhood, 24 Aug 2005. MZUSP 90285, 28, 12.7 -21.0 mm SL, S��o Paulo, Embu, stream affluent of rio Embu, 25 Mar 2006. MZUSP 90292, 31 (10, 2 c&s, 17.5-26.1 mm SL), 17.5-26.1 mm SL, Embu, stream affluent of rio Embu-Mirim, near deposit of construction material Lagoa, on road Jo��o Rodrigues de Morais, 25 Mar 2006. MZUSP 90309, 14, 11.4-22.3 mm SL, S��o Paulo, stream affluent of Guarapiranga dam, on road of Jaceguava, near soccer stadium, 26 Mar 2006. MZUSP 95314, 34, 16.8-22.7 mm SL, Embu, ribeir��o Embu (lowland), 29 Aug 2007. MZUSP 99700, 54, 14.6-28.7 mm SL, Embu, stream affluent of Ribeir��o Embu- Mirim, stake 34101 of lot 5 Rodoanel M��rio Covas (upstream), 21 May 2008. MZUSP 101342, 2, 20.4-20.7 mm SL, Itapecerica da Serra, ribeir��o Itaquaxiara, affluent of rio Embu-Mirim, near road of Pedreira, Lagoa neiborhood, 18 Nov 2008. MZUSP 101345, 46, 14.8-21.9 mm SL, Embu, stream affluent of rio Embu-Mirim, stake 34101 of lot 5 Rodoanel M��rio Covas, 02 Feb 2009. MZUSP 102799, 3, 19.9-21.7 mm SL, Biritiba Mirim, in the area of VCN Mining, 13 Mar 2009. MZUSP 102803, 5, 17.0- 17.8 mm SL, S��o Paulo, Biritiba Mirim, Tiet��, rio Tiet��, in the area of VCN Mining, 13 Mar 2009. MZUSP 109043, 26, 11.2-16.5 mm SL, Itapecerica da Serra, ribeir��o Itaquaxiara, affluent of rio Embu-Mirim, near road of Pedreira, Lagoa neighborhood, 21 Aug 2009. MZUSP 109046, 2, 18.4 -23.0 mm SL, Itapecerica da Serra, ribeir��o Itaquaxiara, affluent of rio Embu-Mirim, near road of Pedreira, Lagoa neighborhood, 06 Nov 2009. MZUSP 109052, 46, 9.9 -21.0 mm SL, Embu, stream affluent of ribeir��o Embu-Mirim, stake 34101 of l, Published as part of Carvalho, Fernando R., de Jesus, Guilherme C. & Langeani, Francisco, 2014, Redescription of Hyphessobrycon flammeus Myers, 1924 (Ostariophysi: Characidae), a threatened species from Brazil, pp. 247-256 in Neotropical Ichthyology 12 (2) on pages 248-254, DOI: 10.1590/1982-0224-20130197, http://zenodo.org/record/4551025, {"references":["Gery, J. 1977. Characoids of the World. T. F. H. Publications, Neptune City, NJ.","Mazzoni, R., C. A. Figueiredo, C. R. S. F. Bizerril, G. W. Nunan, K. Tanizaki-Fonseca, N. A. Menezes, O. C. M. Filho & P. A. Buckup. 2000. Peixes. Pp. 63 - 73. In: Bergallo, H. G., C. F. D. Rocha, M. A. S. Alves & M. V. Sluys (Orgs.). A Fauna Ameacada de extincao do Estado do Rio de Janeiro. Ed. UERJ, FAPERJ.","Marceniuk, A. P., A. W. S. Hilsdorf & F. Langeani. 2011. The ichthyofauna from the headwaters of the rio Tiete, Sao Paulo, Brazil. Biota Neotropica, 11: 217 - 236.","Diario Oficial da Uniao. 2004. Lista Nacional das Especies de Invertebrados Aquaticos e Peixes Ameacados de Extincao. Secao 1, n ° 102, 28 de maio de 2004. Available from http: // www. biodiversitas. org. br / f _ ameaca / anexo 1. pdf (22 Oct 2013)","Froese, R. & D. Pauly (Eds.). 2013. FishBase. World Wide Web electronic publication, version (08 / 2013). Available from: http: // www. fishbase. org (16 September 2013)."]}

Moenkhausia doceana (Steindachner, 1877) Figs. 1-5 Tetragonopterus doceanus Steindachner, 1877: 572-575. Type locality: rio Doce. Moenkhausia doceana (Steindachner): - Eigenmann, 1910: 437 (listed). - Eigenmann, 1917: 66; 73-75 (short redescription of species, but without analysis of syntypes). -Zarske & G��ry, 2001: 5-13 (comparison to M. margitae and data of some syntypes). -Zarske & G��ry, 2002: 52 (comparison to M. dorsinuda). - Lima et al., 2003: 148 (listed). -Bertaco & Lucinda, 2006: 65, 66 (comparison to M. pankilopteryx). - Lima et al., 2007: 58 (listed). Diagnosis. Moenkhausia doceana is promptly distinguished from all congeners [except M. affinis (Steindachner), M. barbouri Eigenmann, M. dorsinuda Zarske & G��ry, M. jamesi Eigenmann, M. justae Eigenmann, M. latissima Eigenmann, M. levidorsa Benine, M. margitae Zarske & G��ry, M. moisae G��ry, Planquette & Le Bail, M. newtoni, M. simulata (Eigenmann), M. tergimacula Lucena & Lucena] by a long anal fin, with 29-34 (mode 32) branched rays (vs. 28 or less in others species). Moenkhausia doceana differs from the aforementioned species, except M. tergimacula, by having 4-7 (mode 5) maxillary teeth (vs. 3 or less). Lastly, M. doceana differs of M. tergimacula by having 7-8 (mode 7) scale rows above lateral line and no marks anteriorly to dorsal-fin origin (vs. 5�� scales rows above lateral line and presence of a distinct dark, saddle-like mark on the body immediately anterior to dorsal-fin origin). Description. Morphometric data are summarized in Table 2. Body compressed, moderately long and high, greatest body depth at vertical through dorsal-fin origin. Dorsal profile of head convex from tip of upper jaw to vertical through anterior nostril; slightly concave or straight from that point to tip of supraoccipital spine. Dorsal profile of body slightly convex from posterior tip of supraoccipital spine to base of first dorsalfin ray, and straight that point to adipose-fin origin. Ventral profile of body convex from tip of lower jaw to pelvic-fin origin, straight or slightly convex from that point to anal-fin origin, and straight and posterodorsally slanted along of analfin base. Dorsal and ventral profile of caudal peduncle straight to slightly concave. Mouth terminal, premaxillary and dentary approximately the same size. Maxilla extending posteroventrally to first onethird of orbit, almost reaching tip of second infraorbital, approximately at 45 degrees angle relative to longitudinal axis of body. Main axis of maxilla straight, with approximately same width along entire length. Nostrils close to each other, anterior opening small and circular, posterior opening twice in size and reniform. Nostrils separated by skin flap when adpressed, almost covering the posterior nostril. Frontals not united anteriorly, with a triangle-shaped fontanel; parietal fontanel large, extending from epiphyseal bar to supraoccipital spine. Eyes relatively large, without adipose eyelid. Infraorbital bones complete, with six elements (Fig. 4). Laterosensory canal from first to sixth infraorbital close to inner margin of orbital rim. Third infraorbital largest, approximately twice the size of others (except second), with inferior margin slightly straight (Fig. 4), not contacting the laterosensory canal of preopercle ventrolaterally. Two tooth rows in the premaxilla: outer row with 4(13), 5(48), or 6(6) [4(4) or 5(2)] tricuspid teeth; inner row with 5(34) or 6(33) [5(6)], pentacuspid teeth. Dentary with 4(42) or 5(14) [4(3) or 5(3)] large, pentacuspid teeth, followed by a series of 10(1), 11(1), or 13(2) conical or triscupid teeth, abruptly smaller than the or five anterior largest teeth. Dorsal border of maxilla relatively straight. Maxilla with 4(12), 5(36), 6(15), or 7(7) [4(1), 5(3), or 6(2)] tri- or tetracuspid teeth along anteroventral margin (Fig. 5). Central median cusp of all teeth longer than remaining lateral cusps; cusps tips slightly curved inwardly on dentary and premaxillary teeth. Scales cycloid, with few radii (2-6), relatively small; circuli marked anteriorly and marginally (dorsal and ventral). Lateral line complete, extending from the superior portion of opercular opening to beginning of caudal fin with 34(2), 35(31), 36(29), 37(2), or 38(1) [34(2), 35(2), or 36(2)] perforated scales; horizontal scales rows between dorsal-fin origin and lateral line 7(23), 7��(10), or 8(37) [7(1) or 8(3)]; horizontal scales rows between lateral line and pelvic-fin origin 6(4), 6��(4), or 7(49) [6(1), 6��(1), or 7(2)]. Predorsal scales 10(20) or 11(13) [10(2) or 11(4)]. Scale sheath along anal-fin base 14(3), 15(2), 16(8), 17(4), 18(2), or 19(2) [15(1) or 16(1)] in a single row. Circumpeduncular scales 14(33) or 15(12) [14(4) or 15(1)]. Axillary scale present, covering two scales posteriorly, distinctly triangular-shaped, thinner on posterior tip. Dorsal fin rays ii,9(77) [ii,9(6)]; first unbranched ray approximately half the length of second one or shorter. Dorsalfin origin slightly forward on midbody, at vertical through anterior one-third of pelvic-fin origin. Base of last dorsal-fin ray at vertical through distal tip of longest pelvic-fin ray, anterior to origin of anal fin. Tip of longest ray of adpressed dorsal fin at vertical through base of first two branched analfin rays. First dorsal-fin pterygiophore inserted between neural spine of eighth and ninth precaudal vertebra. Adipose fin present, well-developed. Pectoral fin with i,11(2), 12(15), 13(51), or 14(3) [i,12(3) or i,14(3)]; longest ray extending to one-third of pelvic-fin origin. Pelvic fin with i,7(77) [i,7(6)] rays. Pelvic-fin origin located anterior to vertical through dorsal-fin origin and with small bony hooks in mature males, absent in females. Tip of longest pelvic-fin ray reaching up to second or third branched ray anal-fin ray when adpressed. Anal-fin rays iv,29(2), 30(1), 31(10), 32(26), 33(23), or 34(6) [31(3), 32(2), or 33(1)]. Caudal fin forked, lobes slightly rounded or pointed, upper lobe slightly smaller than lower lobe, with i,9/8,i(77) [i,9/8,i(6)] rays. Caudal fin scaled, scales over base and along lower caudal-fin lobe up to middle; upper lobe with few scales, extending to its anterior one-third; scales gradually decreasing posteriorly in size. Dorsal procurrent caudal-fin rays 9(1), 10(1), or 11(2), and ventral procurrent caudal-fin rays 7(2), or 8(2). Branchiostegal rays 4(4). First gill arch with 8(4) gill rakers on epibranchial, 1(4) between epibranchial and ceratobranchial, 10(3) or 11(1) on ceratobranchial, and 2(4) on hypobranchial. Denticles on gill rakers present. Precaudal vertebrae 15(4) and caudal vertebrae 19(2) or 20(2). Total vertebrae 34(2) or 35(2). Supraneurals 3(1) or 4(3), filiform, with dorsal portion expanded and some bifurcated dorsally. Color in alcohol. Overall ground coloration yellowish, with light brown thin stripe extending longitudinally along lateral line from vertical through dorsal-fin origin to caudal peduncle. Head brownish dorsally, paler towards sides of head. Dark chromatophores concentrated on distal margin of scales resulting in reticulated pattern. Mid-dorsal region darker than flanks. Humeral region with horizontally rounded dark spot immediately above lateral-line scales, extending about two scales vertically and 4-6 scales horizontally. A second faint blotch behind humeral spot, comma shaped. Area above hypural plate with faint dark spot, almost triangle shaped. First rays of pelvic and anal fins with dark chromatophores, resulting in darker fin border. Dorsal fin with scattered dark chromatophores, more concentrated on anterior one-half.Anal fin with scattered dark chromatophores.Adipose-fin contour with dark chromatophores. Remaining fins hyaline on tips, with scattered dark chromatophores between unbranched rays (Fig. 2). Color in life. The following description is based on the examination of freshly specimens recently caught along northern Esp��rito Santo and southern Bahia rivers. Ground color pale yellow to silver gray, darker dorsally. Sides of body somewhat silver colored anteriorly to light red posteriorly. Longitudinal inconspicuous brilliant stripe at level of lateral line, silvery colored in some specimens. Humeral region with a faint horizontally rounded spot, followed by a second comma shaped faint blotch. Eyes yellowish to whitish. Opercle and preopercle region silver colored; fins yellowish to translucent (Fig. 3). A faint brownish spot, almost triangle shaped on caudal peduncle. Sexual dimorphism. Bony hooks on anal and pelvic fins, and also anal-fin profile distinguish males from females in Moenkhausia doceana. Mature males with very small bony hooks on the segments of the last unbranched and all branched anal-fin rays (one pair of bony hooks on posterior surface of hemitrichia), absents in females. Males with anal-fin distal margin slightly straight (Fig. 2a) while females with anal-fin distal margin falcate anteriorly (Fig. 2b). Gill glands (sensu Burns & Weitzman, 1996) were not found macroscopically on first gill arch on both sexes. Distribution. Moenkhausia doceana occurs along coastal river systems, with northern limits corresponding to the rio Jo��o de Tiba, on extreme southern Bahia river basins. The species is recorded until southern limits at the rio Riacho, a small coastal basin south of rio Doce in Esp��rito Santo State, and to the West in the middle rio Doce lake systems, rio Doce basin (Fig. 6). Known populations of M. doceana inhabit areas of mild relief, as in lakes of the rio Doce valley and along coastal drainages between Esp��rito Santo and southern Bahia states. The western portion of its distribution is within Coastal Tablelands area, a wide region with elevations usually not passing 150 m, coinciding with the Cenozoic sediments of the Barreiras Group, well seen along the whole coastal distributional area. Despite its range distribution, no significant morphological and meristic differences were found among M. doceana populations, suggesting that the species corresponds to a single taxon in all coastal drainages where it occurs. Regarding global biogeography regionalization of freshwater systems, the pattern of distribution of M. doceana fits the Northeastern Mata Atl��ntica ecoregion (sensu Abell et al., 2008). Gerais (Ottoni et al., 2011), Reserva Biol��gica do C��rrego Grande, Esp��rito Santo (Sarmento-Soares & Martins-Pinheiro, 2013), and Reserva Biol��gica Sooretama, Esp��rito Santo (LMSS and RFMP, pers. obs.). Popular name. Moenkhausia doceana is known as lambari (Lima et al., 2003) or piaba, in northern Esp��rito Santo and southern Bahia states. Ecological notes. Moenkhausia doceana inhabits second and third order streams, about two to three meters wide and 0.5 to 1.6 meters deep, characterized by clear water and moderate to fast current, sandy bottom with gravel, and riparian surrounding vegetation. The fishes were captured in the water column, mainly near the surface. It is an invertivore that eats aquatic insect larvae (e.g., Chironomidae), nymphs (e.g., Ephemeroptera), and fragments of terrestrial arthropods (ants, beetles, and spiders). Moenkhausia doceana occurs syntopically with Characidium aff. fasciatum Reinhardt, Hoplias malabaricus (Bloch), Astyanax aff. intermedius Eigenmann, Astyanax aff. lacustris (L��tken), Hyphessobrycon bifasciatus Ellis, Trichomycterus pradensis Sarmento-Soares, Martins-Pinheiro, Aranda & Chamon, Corydoras nattereri Steindachner, Pimelodella aff. vittata (L��tken), Poecilia vivipara Bloch & Schneider, and Geophagus brasiliensis (Quoy & Gaimard) (Sarmento-Soares & Martins Pinheiro, 2013). Moenkhausia doceana was recorded in conservation units such as the Parque Estadual do Rio Doce (PERD), Minas Phylogenetic reconstruction. Phylogenetic analysis, according to implied weighting procedures (Mirande, 2009, 2010), with consensus of ���k��� = 10.07571802 (three trees) and ���k��� = 11.06353351 (three trees) values, resulted in one cladogram with 2263 steps, Consistency Index (CI) of 0.16 and Retention Index (RI) of 0.66. Moenkhausia doceana is included in the Tetragonopterinae of Mirande (2010). It is sister group of (Tetragonopterus argenteus (Gymnocorymbus ternetzi (Stichonodon insignis (Poptella paraguayensis, Stethaprion erythrurus)))). Synapomorphies for this clade are: ch. 201:1 - denticles on gill rakers absent [reversed in M. doceana]; ch. 287:1 - 18 or more branched anal-fin rays; ch. 314:0 - bony hooks on last pelvic-fin ray of adult males absent; ch. 341:1 - humeral spot horizontally-ovate. Autapomorphies for M. doceana are: ch. 22:0 - frontal fontanel present; ch. 63:0 - anterior region of third infraorbital not much expanded relative to posterior region of second infraorbital; ch. 66:0 - fourth infraorbital present and well developed; ch. 82:0 - dorsal end Syntypes Non-type specimensRange Mean Range Mean SD N Standard Length (mm) 57.8-83.8 71.1 30.1-81.4 54.8 - 73Percents of Standard LengthBody depth 41.0-45.7 44.0 37.0-45.8 41.9 2.0 73 Head length 24.6-27.3 26.1 16.6-30.9 26.7 1.9 73 Head depth 31.6-35.7 33.6 27.0-35.7 32.3 2.6 56 Predorsal length 51.3-52.6 52.0 48.9-54.4 51.0 1.3 73 Prepelvic length 42.1-45.0 44.1 40.9-48.5 44.6 1.5 73 Pelvic fin to anal distance 16.2-18.6 17.0 13.3-21.5 16.3 1.4 70 Caudal peduncle depth 10.3-11.6 10.7 7.8-11.8 9.8 0.8 73 Dorsal-fin base length 13.6-14.3 13.9 12.4-17.1 14.2 0.8 73 Anal-fin base length 41.0-44.0 42.2 37.4-44.2 41.0 1.5 73 Pectoral-fin length 20.5-24.0 22.4 21.0-25.5 23.0 0.9 72 Pelvic-fin length 17.7-20.3 19.2 16.3-22.1 19.2 1.0 72 Dorsal-fin length 29.5-34.2 31.9 28.4-35.2 32.2 1.7 71 Anal-fin length 20.0-23.3 21.7 18.4-25.3 22.2 2.0 61 Caudal peduncle length 9.9-11.7 10.8 9.3-13.4 11.0 0.9 73 Dorsal fin to adipose-fin distance 40.1-42.6 41.0 37.1-42.7 40.0 1.1 73 Eye to dorsal-fin origin 36.1-39.3 37.9 33.9-40.1 36.4 1.4 73 Dorsal origin to caudal origin 55.2-58.1 56.5 50.3-59.8 56.4 1.4 73Percents of head lengthInterorbital width 32.5-35.3 33.9 27.5-36.1 31.3 1.7 70 Snout length 20.8-24.9 23.1 21.9-29.3 24.8 1.9 65 Orbital diameter 39.9-44.0 42.3 40.6-48.5 44.6 1.9 55 Upper jaw length 45.7-48.5 47.7 41.3-49.8 47.3 1.9 41 of laterosensory canal of preopercle and suprapreopercle not overlapping anterodorsal process of opercle; ch. 109:1 - ventral margin of horizontal process of anguloarticular perpendicular to laterosensory canal of dentary from medial view; ch. 135:1 - two or more maxillary teeth; ch. 225:0 - abdominal ribs on anterior caudal vertebrae absent; ch. 281:0 - seven or fewer supraneurals; ch. 305:0 - anterior ventral procurrent caudal-fin rays paired, only distally fused; ch. 315:1 - bony hooks on first pelvic-fin ray of adult males present; ch. 340:1 - insertion of pterotic aponeurosis on a lobe situated dorsal to horizontal semicircular canal. Material examined. Syntypes. All from Brazil, rio Doce. Tetragonopterus doceanus, NMW 57389, 2(2), 57.8-70.0 mm SL. NMW 57591, 2(2), 83.3- 83.3 mm SL. MNHN 1913-135, 1(1), 68.4 mm SL. MHNH 1913-136, 1(1), 62.2 mm SL. Moenkhausia doceana, Non-type specimens (ordered by river basin, and munic��pio). All from Brazil. Esp��rito Santo State, rio Riacho basin. Munic��pio de Aracruz. MBML 2728, 4, 57.4-65.0 mm SL (ex-CIUFES 130078), limit between Aracruz and Linhares, Lagoa do Aguiar. Esp��rito Santo State, rio Doce basin. Munic��pio de Colatina. MBML 678, 2(1), 21.7-30.1 mm SL, mouth of rio Santa Joana. MBML 3391, 1, 34.3 mm SL, rio Santa Maria do Rio Doce. MBML 3590, 1(1), 61.0 mm SL, c��rrego Miracema on right side of road ES-080 from S��o Domingos do Norte to Colatina. MNRJ 37747, 1, 56.2 mm SL, same data as MBML 3590. Munic��pio de Itarana. MBML 763, 1, 26.4 mm SL, rio Santa Joana. Munic��pio de Linhares. MCP 42079, 1, 43.6 mm SL, rio Pequeno. MNRJ 9708, 1, not measured, Lagoa Juparan��. MZUSP 1461, 2, 74.6-80.0 mm SL, rio Doce. MZUSP 1560, 6, 52.6-63.4 mm SL, rio Doce. MZUSP 1622, 3, 47.7-75.7 mm SL, rio Doce. MZUSP 17065, 1, 53.0 mm SL, Lagoa do Aviso. MZUSP 17069, 1, 42.6 mm SL, Lagoa Nova. MZUSP 17073, 1, 75.4 mm SL, Lagoa Juparan��. MZUSP 75137, 11, 52.4-75.9 mm SL, rio Doce. ZUEC 3583, 4, 18.2-25.6 mm SL, c��rrego Rancho Alto, Reserva Florestal Vale do Rio Doce. Munic��pio de Pancas. MBML 1749, 2(1), 35.7-36.6 mm SL, c��rrego Ub��, tributary of rio Pancas. MBML 1836, 2(1), 49.6- 22.9 mm SL, c��rrego Floresta, tributary of rio Pancas on fazenda Floresta Lajinha. MBML 2330, 2(2), 40.0- 49.8 mm SL, c��rrego S��o Bento, tributary of rio Pancas in S��o Bento. Munic��pio de Santa Teresa. MZUSP 1562, 1, 64.2 mm SL, rio 25 de Julho. MZUSP 1629, 3, 57.6-62.8 mm SL, rio 25 de Julho. Minas Gerais State, rio Doce basin. Munic��pio de Frei Inoc��ncio. MCP 17771, 3, 30.5-30.9 mm SL, rio Sua��u�� tributary of rio Doce, under bridge on road BR-116. Munic��pio de Maril��ria. MZUSP 28972, 1, 58.9 mm SL. Lago Dom Helv��cio, rio Doce Valley. MZUSP 28981, 15, 55.7-66.7 mm SL, Lagoa Carioca, rio Doce Valley. MZUSP 36648, 2, 57.1-67.8 mm SL, Lagoa Carioca, rio Doce Valley. MZUSP 107342, 5, 61.1-72.7 mm SL, Lagoa Carioca, Parque Estadual Rio Doce. Esp��rito Santo State, rio Barra Seca basin. Munic��pio de Linhares. MBML 48, 5(3), 41.5- 60.4 mm SL, rio Ipiranga in Pontal do Ipiranga. MCP 17769, 4, 20.1-48.0 mm SL, Lagoa do Cupido on road BR-101, on northern limits of Reserva Biol��gica de Sooretama. MCP 17770, 1, 32.5 mm SL, coastal river on km 118 of road BR-101, near Reserva Biol��gica de Sooretama. Munic��pio de Sooretama. MBML 5114, 52(7), 59.2-81.4 mm SL, second pool on road ES-356 near bridge over rio Barra Seca, inside Reserva Biol��gica de Sooretama. MBML 5169, 5(2), 35.4-42.2 mm SL, rio Barra Seca under bridge on road ES-356 around the area of Reserva Biol��gica de Sooretama. MNRJ 5342, 7, 55.1-64.0 mm SL, rio Barra Seca, on road Linhares to S��o Mateus (in precary conservation state). MNRJ 17588, 22, 38.6- 64.6 mm SL, rio Barra Seca. MNRJ 40099, 10, 46.2-76.5 mm SL, c��rrego Cupido, near road BR-101 inside Reserva Biol��gica de Sooretama. MNRJ 40103, 1, 44.1 mm SL, c��rrego Parais��polis on its mouth in rio Barra Seca inside Reserva Biol��gica de Sooretama. MNRJ 40115, 6, 26.4-43.7 mm SL, rio Barra Seca under bridge on road ES-356 around the area of Reserva Biol��gica de Sooretama. MNRJ 40259, 54, 29.1-63.4 mm SL, same data as MBML 5169. Munic��pio de Jaguar��. MBML 5186, 1(1), 68.0 mm SL, rio Barra Seca on road between Nossa Senhora de F��tima and Vila Val��rio, around the area of Reserva Biol��gica de Sooretama. MNRJ 5324, 1, not measured, rio Cachimbau, on road Linhares to S��o Mateus. MNRJ 10916, 19, not measured, same data as MNRJ 5324. MN, Published as part of Carvalho, Fernando R., Sarmento-Soares, Luisa M. & Martins-Pinheiro, Ronaldo F., 2014, Redescription of Moenkhausia doceana (Steindachner, 1877) (Ostariophysi: Characiformes): a characid from the Northeastern Mata Atl��ntica ecoregion, Brazil, pp. 377-388 in Neotropical Ichthyology 12 (2) on pages 378-383, DOI: 10.1590/1982-0224-20140006, http://zenodo.org/record/4639092, {"references":["Steindachner, F. 1877. Die Susswasserfische des sudostlichen Brasilien (III). Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Classe, 74: 559 - 694.","Eigenmann, C. H. 1910. Catalogue of the fresh-water fishes of tropical and south temperate America. In: Scott, W. B. (Ed.). Reports of the Princeton University Expeditions to Patagonia 1896 - 1899. Princeton, N. J., 3: 375 - 511.","Eigenmann, C. H. 1917. The American Characidae - I. Memories of the Museum of Comparative Zoology, 43: 1 - 102.","Lima, F. C. T., L. R. Malabarba, P. A. Buckup, J. F. Pezzi da Silva, R. P. Vari, A. Harold, R. Benine, O. T. Oyakawa, C. S. Pavanelli, N. A. Menezes, C. A. S. Lucena, M. C. S. L. Malabarba, Z. M. S. Lucena, R. E. Reis, F. Langeani, L. Casatti, V. A. Bertaco, C. Moreira & P. H. F. Lucinda. 2003. Genera Incertae Sedis in Characidae. Pp. 106 - 169. In: Reis, R. E., S. O. Kullander & C. J. Ferraris Jr. (Eds.). Check List of the freshwater fishes of South and Central America. Porto Alegre, Edipucrs.","Lima, F. C. T., P. A. Buckup, N. A. Menezes, C. A. S. Lucena, Z. M. S. Lucena, M. Toledo-Piza & A. Zanata. 2007. Familia Characidae: generos incertae sedis. Pp. 44 - 62. In: Buckup, P. A., N. A. Menezes & M. S. Ghazi. Catalogo das especies de peixes de agua doce do Brasil. Serie livros 23, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro.","Mirande, J. M. 2010. Phylogeny of the family Characidae (Teleostei: Characiformes): from characters to taxonomy. Neotropical Ichthyology, 8: 385 - 568.","Abell, R., M. L. Thieme, C. Revenga, M. Bryer, M. Kottelat, N. Bogutskaya, B. Coad, N. Mandrak, S. L. Balderas, W. Bussing, M. L. J. Stiassny, P. Skelton, G. R. Allen, P. Unmack, A. Naseka, R. Ng, N. Sindorf, J. Robertson, E. Armijo, J. Y. Higgins, T. J. Heibel, E. Wikramanayake, D. Olson, H. L. Lopez, R. E. Reis, J. G. Lundberg, M. H. S. Perez & P. Petry. 2008. Freshwater ecoregions of the world: a new map of biogeographic units for freshwater biodiversity conservation. Bioscience, 58: 403 - 414.","Ottoni, F. P., A. Q. Lezama, M. L. Triques, E. N. Fragoso-Moura, C. C. T. Lucas & F. A. R. Barbosa. 2011. Australoheros perdi, new species (Teleostei: Labroidei: Cichlidae) from the lacustrine region of the Doce River Valley, southeastern Brazil, with biological information. Vertebrate Zoology, 61: 137 - 145.","Sarmento-Soares, L. M. & R. F. Martins Pinheiro. 2013. A fauna de peixes na REBIO Corrego Grande e seu entorno direto, Espirito Santo, Brasil. Boletim do Museu de Biologia Mello Leitao, 31: 25 - 57.","Mirande, J. M. 2009. Weighted parsimony phylogeny of the family Characidae (Teleostei: Characiformes). Cladistics, 2009: 573 - 613."]}

Ectrepopterus uruguayensis Fowler 1943 (Figs. 1���6; Table 2) Megalamphodus uruguayensis Fowler 1943: 313 (type locality: Uruguay, no further locality data available; type species of the subgenus Ectrepopterus by original designation and monotypy; mistyped also as Magalamphodus uruguayensis). Megalamphodus (Ectrepopterus) uruguayensis. G��ry 1972: 14 (redescription of holotype). Megalamphodus uruguayensis. G��ry 1977: 586 (in key). Hyphessobrycon uruguayensis. Weitzman and Palmer 1997: 234 (listed as a valid species probably belonging to the rosy tetra group). Thomaz et al. 2010 (phylogeny based on DNA sequences). Diagnosis. The same for the genus. The larger number of maxillary teeth (6���11) and the presence of longitudinal, wavy stripes laterally on body in larger specimens further distinguish this species from several characids. Description. Morphometric data are summarized in Table 2. Body compressed and elongate; greatest body depth near to dorsal-fin origin. Dorsal head profile nearly straight. Dorsal body profile convex from supraoccipital bone to base of last dorsal-fin ray and straight from this point to adipose-fin origin. Ventral profile of head smoothly convex. Ventral body profile slightly convex to nearly straight from pectoral-fin origin to pelvic-fin origin, and straight to anal-fin origin. Body profile along anal-fin base posterodorsally slanted. Caudal peduncle elongate, nearly straight to slightly concave along dorsal and ventral margins. tion. The range not includes the holotype. Mouth terminal, lower jaw projecting slightly more than upper jaw. Maxilla long and slightly curved aligned at angle of approximately 45 degrees relative to longitudinal body axis. Maxilla slightly widened anteroposteriorly, and extends to junction of infraorbital 2 and 3. Premaxilla with one or two tooth rows: outer row, when present, with one to two tricuspid teeth; inner row with six to seven tricuspid teeth (7 *), gradually decreasing in length from first to fourth or fifth teeth and last two smaller. Maxilla with six to eleven (8 *, mode = 10) tricuspidate teeth; and last five or six uni- to tricuspid teeth. Four or five anteriormost dentary teeth larger, tricuspid, followed by one or two medium sized tricuspid teeth, and eight to 13 teeth with one to three cusps. Central cusp in all teeth two to three times longer and broader than other cusps. All cusp tips slightly curved posteriorly towards oral cavity (Fig. 4). Dorsal-fin rays ii, 8���10 (ii, 9 *; n = 93; ii, 8 and ii, 10 in one specimen each); first unbranched ray approximately one-half length of second ray. Dorsal-fin origin located at middle of SL and posterior to vertical through pelvic-fin origin. Distal margin of dorsal fin slightly convex. Adipose-fin located approximately at vertical through insertion of last anal-fin ray. Pectoral-fin rays i, 10���12 * (mode = 10, n = 93). Pelvic-fin rays i, 6 or 7 * (mode = 7, n = 93). Pelvic-fin origin located anterior to vertical through dorsal-fin origin. Anal-fin rays iii���v, 19���26 (22 *, rarely 19 or 24���26, mode = 22, n = 93). First unbranched ray normally only apparent in cleared and stained specimens. Anal-fin origin posterior to vertical through base of last two dorsal-fin ray. Caudal-fin forked, lobes similar in size, with 18��� 20 principal rays (19 *, two with 18 and one with 20, n = 92). Dorsal procurrent rays 9���11 (mode = 10, n = 10), and ventral procurrent rays 8���10 (mode = 9, n = 10). Scales cycloid, moderately large. Lateral line incomplete, perforated scales 5���9 (8 *, one specimen with 5 and one specimen with 9, mode = 6, n = 93). Longitudinal scale series including pored scales 32 *��� 36 (mode = 33, n = 93). Scale rows between dorsal-fin origin and lateral line 6 or 7 * (mode = 6, n = 93); scale rows between lateral line and pelvic-fin origin 4���6 (5 *, mode = 5, n = 93). Predorsal scales 10 *��� 13, arranged in regular series (mode = 11, n = 82). Scale rows around caudal peduncle 14 * (n = 93). Scale sheath along anal-fin base 6���12 (8 *, mode = 8) scales in single series, extending to base of fifth to tenth branched rays. Precaudal vertebrae 16 * or 17 (mode = 16, n = 11); caudal vertebrae 16 or 17 * (mode = 17, n = 11); total vertebrae 33 * or 34 (mode = 33, n = 11). Supraneurals 5 or 6 (mode = 6, n = 10). Gill rakers on upper limb of outer gill arch 5 or 6 (mode = 6), and on lower limb 9���11 (mode = 9, n = 10). Ventral extent of third infraorbital reaching horizontal arm of preopercle, but not reaching the laterosensory canal of preopercle. Fontanels forming slightly narrow groove; parietal little longer than frontal. Frontals not contacting anteriorly to frontal fontanel. Color in alcohol. The holotype is discolored, except for the humeral spot (Fig. 1). Color description based on non-type specimens (Fig. 5). Dorsal and dorsolateral portion of head and body pigmented dark brown. Scales on lateral and dorsal surface of body with scattered dark brown chromatophores. In some specimens larger than 35.0 mm SL the dorsal and ventral border of scales are dark brown pigmented forming a wavy, striped pattern between longitudinal rows of scales, more conspicuous in middle portion of body (Fig. 5). One narrow humeral spot vertically elongate, located over third to fourth lateral line scales and extending over two longitudinal series of scales above and below of lateral line. Midlateral body stripe very narrow extending from posterior middle region of body to base of median caudal-fin rays, wide and more densely pigmented on caudal peduncle forming a triangle spot. Anteriorly directed chevron-shaped marks along the midlateral body. Small black chromatophores scattered over rays of all fins. Caudal-fin rays darkened black in the distal portion. Adipose-fin smoothly dark black pigmented along the distal border. Color in life. Color pattern similar to described for alcohol preserved specimens. Overall body and head color pattern silvery to golden (Figs. 5 and 6). Chromatophores distribution pattern of head, body and fins as described above for alcohol preserved specimens. Eye with upper third of iris iridescent red. Dorsal fin yellowish. Pectoral fin hyaline. Pelvic fin orangish. Anal fin yellowish, more intense along proximal half of most anterior anal-fin rays. Adipose fin yellowish. Caudal fin yellowish, mostly at the proximal half of the dorsal and ventral caudal-fin lobes; middle caudal-fin rays and distal third portion or distal half of caudal-fin rays with dark black melanophores (Figs. 5 and 6). Wavy, striped pigmentation pattern along the scales is not clearly discernible in living specimens. Sexual dimorphism. Mature males of Ectrepopterus uruguayensis are recognized by the presence of small bony hooks on all rayed fins (absent in females). Mature males present bony hooks in the third distal portion of the first to fifth branched rays, and in the distal portion of all pectoral-fin rays. Pelvic fin usually bearing bony hooks along ventromedial border of the unbranched and first to fourth branched rays. Anal-fin rays bearing one small, elongate, retrorse bony hook along posterolateral border of each segment of lepidotrichia, usually along last unbranched ray to last branched ray, in the distal portion of all caudal-fin rays, and in the two posteriormost procurrent caudal-fin-rays. Hooks usually located along of distal portion of each ray. Anal-fin profile nearly straight in males and smoothly concave in females. The presence of bony hooks is often found in males, but only mature males (specimens larger than 33.0 mm SL) possess bony hooks on all rayed fins. Pectoral-fin tip reaching pelvic-fin origin in males but not in females. Pelvic-fin tip surpass anal-fin origin in males but not in females. Males and females also differ by anal-fin profile, which is nearly straight in males and smoothly concave in females. Gill glands were not found on first gill arch on both mature males and females. Distribution. Ectrepopterus uruguayensis occurs in the lower tributaries of the r��o Uruguay (r��o Cuareim, r��o Salto, and r��o Negro basins), and r��o de La Plata (r��o Santa Luc��a and arroyo Pand), in Uruguay (Fig. 7). The Uruguay River basin in Brazil has been extensively sampled for the last 20 years, with intensive collecting undertaken in some localities, and so far there are no records of the species in this country. Menni (2004) listed Hyphessobrycon uruguayensis (= E. uruguayensis) for Argentina without any specific location or list of examined material. Ecological notes. Ectrepopterus uruguayensis inhabits streams, and occurs in semi-lentic and lotic shallow areas (up to 1.2 m deep) with moderate submerged vegetation and riparian vegetation composed by trees and shrubs or absent. The streams have transparent water, and bottom with mud, some stretches with sand or gravel. Water temperature measured during field work ranged from 10 to 30 ��C, depending on the season. The pH was found around 7 in the neutral range. Keeping these conditions under consideration, Ectrepopterus uruguayensis can be easily kept in aquaria up to eight years. This species is usually collected in small numbers in natural habitats. Observations in aquaria (TOL) showed that, contrary to many other characids, it does not form schools, but rather specimens stay mainly alone, without showing territorial behavior. Material examined. All from Uruguay: ANSP 70331, holotype, female, 29.2 mm SL, 1935, F. Felippone. MCP 13080, 5 (1 c&s), 27.7���36.5 mm SL, arroyo Catal��n, puente de la ruta 30, Artigas, ca. 30 �� 50 ���S 56 �� 14 ���W, 9 Jul 1987, L. H. Amato. MCP 31907, 3 (1 c&s), 33.3���39.7 mm SL, arroyo Salsipuedes, tributary of the r��o Tacuaremb��, Tacuaremb��, 32 �� 20 ��� 39 ������S 56 �� 15 ��� 13 ������W, 15 Aug 2002, P. Laurino et al. MCP 31915, 35.9 mm SL, arroyo Catal��n Grande, Artigas, 30 �� 50 ��� 40 ������S 56 �� 14 ��� 30 ������W, 16 Aug 2002, P. Laurino et al. MCP 33504, 2, 30.2��� 35.9 mm SL, arroyo Saucedo, Salto, 31 ��06��� 28 ������S 57 �� 30 ��� 25 ������W, 17 Mar 2003, P. Laurino et al. MCP 33505, 6 (2 c&s), 23.6���29.1 mm SL arroyo Palomas (loc 2), Salto, 31 ��04��� 43 ������S 57 �� 37 ��� 26 ������W, 17 Mar 2003, P. Laurino et al. MCP 33506, 4, 27.1���28.7 mm SL, arroyo Palomas (loc 1), Salto, ca. 31 ��03��� 16 ������S 57 �� 37 ��� 43 ������W, 17 Mar 2003, P. Laurino et al. MCP 33507, 4, 25.5���33.3 mm SL, arroyo Cardoso, San Jos��, 34 �� 24 ��� 50 ������S 56 �� 26 ��� 49 ������W, 23 Mar 2003, E. Lartigau et al. MCP 36790, 33.8 mm SL, arroyo Pando, Canelones, 34 �� 44 ��� 19 ���S 54 �� 56 ��� 27 ���W, 26 Aug 2004, P. Laurino et al. MCP 36791, 2, 38.7���42.1 mm SL, creek near Paso de San Borja, Durazno, 33 �� 24 ��� 57 ���S 56 �� 25 ��� 57 ���W, 22 Aug 2004, P. Laurino et al. MCP 36792, 35.6 mm SL, arroyo Pando, Canelones, 34 �� 42 ��� 12 ���S 55 �� 56 ��� 43 ���W, 31 Aug 2004, T. O. Litz et al. MCP 36795, 13, 29.2���43.4 mm SL, ditch between arroyo Salsipuedes and ruta 5 Km 289.5, tributary of the rio Tacuaremb��, Tacuaremb��, 32 �� 29 ��� 13 ���S 56 �� 22 ��� 44 ���W, 22 Aug 2004, P. Laurino et al. MCP 36796, 8, 28.8���35.8 mm SL, arroyo Catal��n Grande, Artigas, 30 �� 50 ��� 40 ���S 56 �� 14 ��� 30 ���W, 23 Aug 2004, P. Laurino et al.; MTD-F 27461 ���27466, 6, 25.9���35.6 mm SL, arroyo Palomas (loc 2), Salto, 31 ��04��� 43 ������S 57 �� 37 ��� 26 ������W, 17 Mar 2003, P. Laurino et al. UFRGS 7169, 4, 29.0���32.0 mm SL, arroyo Mestre de Campo, on road to Polanco of YI, drainage of r��o YI, Durazno, 33 �� 24 ��� 55 ������S 56 �� 12 ���06������W, 29 May 2005, L. R. Malabarba et al. UFRGS 7170, 11, 30.2 ���40.0 mm SL, arroyo affluent to arroyo Mestre de Campo, on road to Polanco of YI, drainage of r��o YI, Durazno, 33 �� 23 ���S 56 �� 13 ���W, 29 May 2005, L. R. Malabarba et al. UFRGS 7171, 23, 36.2���44.5 mm SL, arroyo on road 26 ca. 59 km of Melo, between arroyos Saule and Fratile Muerto, Melo, 32 �� 17 ��� 39 ������S 54 �� 44 ��� 59 ������W, 28 May 2005, L. R. Malabarba et al. UFRGS 7172, 2, 36.4���39.6 mm SL, marginal pool and arroyo Corrales, on road 27, drainage of r��o Tacuaremb��, Rivera, 31 �� 23 ��� 26 ������S 55 �� 15 ��� 14 ������W, 27 May 2005, L. R. Malabarba et al. UFRGS 7343, 1, 36.7 mm SL, r��o Caraguat��, r��o Tacuaremb�� basin, Tacuaremb��, 32 ��09��� 29 ���S 55 ��01��� 27 ���W, 28 May 2005, L. R. Malabarba et al. UFRGS 7474, 30.6 mm SL, arroyo Cu��apiru, r��o Tacuaremb�� basin, km 12.3 road 27, Rivera, Rivera, 3102 ��� 21 ���S 5529 ��� 31 ���W, 27 May 2005, L. R. Malabarba et al. UFRGS 7506, 47.9 mm SL; UFRGS 10564, 2 ethyl alcohol anhydrous, 38.2 ��� 40.0 mm SL, arroyo on road 26 about 59 km of Melo, Cerro Largo, 32 �� 17 ��� 39 ���S 54 �� 44 ��� 59 ���W, 28 May 2005, L. R. Malabarba et al. UFRGS 7767, 3, 37.2���40.5 mm SL, arroyo Cuar�� Grande on road 4, tributary of r��o Quara��, Artigas, Artigas, 30 �� 47 ���03���S 56 �� 46 ��� 54 ���W, 8 Sep 2005, L. R. Malabarba et al. UFRGS 7785, 1, 37.1 mm SL; UFRGS 12345, 1 ethyl alcohol anhydrous, 38.2 mm SL, arroyo Carpinchuri, tributary of r��o Uruguay on road 3, Paysandu, Paysandu, 31 �� 40 ��� 38 ���S 57 �� 53 ��� 30 ���W, 20 Sep 2005, L. R. Malabarba et al. UFRGS 7899, 1, 39.7 mm SL, arroyo Chapicuy Chico, tributary of r��o Uruguay on road 3, Paysandu, Paysandu, 31 �� 37 ��� 20 ���S 57 �� 52 ��� 51 ���W, 10 Sep 2005, L. R. Malabarba et al. UFRGS 7909, 9, 33.1���40.2 mm SL, ca��ada Cecilia on road 3, km 512, Salto, Salto, 31 �� 16 ���01���S 57 �� 46 ��� 42 ���W, 9 Sep 2005, L. R. Malabarba et al. UFRGS 7918, 2, 38.9���44.4 mm SL, ca��ada on road 4, tributary of r��o Queguay Grande, Paysandu, Paysandu, 32 �� 12 ��� 25 ���S 57 �� 12 ��� 47 ���W, 10 Sep 2005, L. R. Malabarba et al. UFRGS 7945, 36.9 mm SL, ca��ada on road 4, tributary of r��o Queguay Chico, Paysandu, Paysandu, 32 ��01��� 57 ���S 57 �� 19 ��� 30 ���W, 10 Sep 2005, L. R. Malabarba et al. UFRGS 7985, 53, 29.8���44.3 mm SL, arroyo Guaviy�� on road 3, tributary of r��o Uruguay, Artigas, Artigas, 30 �� 37 ��� 51 ���S 57 �� 41 ��� 18 ���W, 9 Sep 2005, L. R. Malabarba et al. UFRGS 8009, 40.2 mm SL, arroyo los Chanchos on road 3, r��o Daym��n basin, Paysandu, Paysandu, 31 �� 28 ��� 34 ���S 57 �� 54 ���06���W, 10 Sep 2005, L. R. Malabarba et al. UFRGS 8073, 9, 33.5���42.9 mm SL, arroyo de las Tunas on road 31, tributary of r��o Arapey Grande, Salto, Salto, 31 �� 20 ���04���S 57 �� 19 ��� 36 ���W, 8 Sep 2005, L. R. Malabarba et al. UFRGS 8105, 2, 42.3���46.4 mm SL, arroyo on road 4, tributary of r��o Valentin Grande, Salto, Salto, 31 �� 16 ��� 32 ���S 57 ��09��� 22 ���W, 8 Sep 2005, L. R. Malabarba et al. UFRGS 8578, 27 (4 c&s), 39.3���40.8 mm SL, arroyo Tala on road 31 between Salto city and Artigas, 31 �� 23 ���09������S 57 �� 33 ��� 46 ������W, L. R. Malabarba et al. UFRGS 10973, ethyl alcohol anhydrous, 32.8 mm SL, arroyo Tala on road 31, tributary of r��o Uruguay, Salto, Salto, 31 �� 23 ���09���S 57 �� 33 ��� 46 ���W, 8 Sep 2005, L. R. Malabarba et al. UFRGS 11129, 2 ethyl alcohol anhydrous, 33.3���38.9 mm SL, arroyo Corrales and lateral puddles on road 27, r��o Tacuaremb�� basin, Rivera, Rivera, 31 �� 23 ��� 26 ���S 55 �� 15 ��� 14 ���W, 27 May 2005, L. R. Malabarba et al. ZVC-P 423, 8, 28.8���38.8 mm SL, arroyo Bol��n, tributary of the arroyo Tom��s Cuadra, r��o Negro drainage, ca. 33 �� 12 ���S 56 �� 10 ���W, E. Messner. ZVC-P 1464, 16 (2 c&s), 31.3���47.4 mm SL, arroyo Higuera, tributary of the arroyo Carpinteria, r��o Negro drainage, Tacuaremb��, ca. 31 �� 45 ���S 55 �� 13 ���W, 5 Oct 1959, E. Messner. ZVC-P 1769, 31.8 mm SL, r��o Santa Luc��a, Arequita, Levalleja, ca. 34 �� 20 ���S 55 �� 15 ���W, Carbonell. ZVC-P 2975, 4, 25.6��� 27.5 mm SL, Estancia San Francisco, 5 km W from Casup��, Florida, ca. 34 ��05���S 55 �� 40 ���W, 12 Apr 1981, E. Lessa & D. Lisandro. ZVC-P 5267, 53, 16.1���24.3 mm SL, arroyo Mahoma Chico, tributary of the r��o San Jos��, San Jos��, ca. 34 ��02���S 56 �� 58 ���W, 4 Jan 1953., Published as part of Malabarba, Luiz R., Bertaco, Vinicius A., Carvalho, Fernando R. & Litz, Thomas O., 2012, Revalidation of the genus Ectrepopterus Fowler (Teleostei: Characiformes), with the redescription of its type species, E. uruguayensis, pp. 47-60 in Zootaxa 3204 on pages 50-56, DOI: 10.5281/zenodo.210066, {"references":["Fowler, H. W. (1943) Notes and descriptions of new or little known fishes from Uruguay. Proceedings of the Academy of Natural Science of Philadelphia, 95, 311 - 334.","Gery, J. (1972) Corrected and supplemented descriptions of certain characoid fishes described by Henry W. Fowler, with revisions of several of their genera. Studies on the Neotropical Fauna, 7, 1 - 35.","Gery, J. (1977) Characoids of the World. T. H. F. Publications, Neptune City, NJ, 672 p.","Weitzman, S. H. & Palmer, L. (1997) A new species Hyphessobrycon (Teleostei: Characidae) from the Neblina region of Venezuela and Brazil, with comments on the putative ' rosy tetra clade'. Ichthyological Exploration of Freshwaters, 7, 209 - 242.","Thomaz, A. T., Malabarba, L. R. & Bonatto, S. L. (2010) The phylogenetic placement of Hollandichthys Eigenmann 1909 (Teleostei: Characidae) and related genera. Molecular Phylogenetics and Evolution, 57, 1347 - 1352.","Menni, R. C. (2004) Peces y ambientes en la Argentina continental. Monografias del Museo Argentino de Ciencias Naturales, No 5, Buenos Aires, 316 p.","Malabarba, L. R. & Weitzman, S. H. (2003) Description of new genus with six new species from southern Brazil, Uruguay and Argentina, with a discussion of a putative characid clade (Teleostei: Characiformes: Characidae). Comunicacoes do Museu de Ciencias e Tecnologia da PUCRS, Serie Zoologia, 16, 67 - 151."]}

Ectrepopterus Fowler 1943 Ectrepopterus Fowler 1943: 313 (subgenus of Megalamphodus; type species by original designation and monotypy: Megalamphodus uruguayensis). G��ry 1972: 14 (redescription of the type species; new diagnosis for the subgenus). G��ry 1977: 586 (synonym of Megalamphodus). Weitzman and Palmer 1997: 234 (synonym of Hyphessobrycon). Diagnosis. The following unambiguous apomorphies obtained through equal weighting analysis diagnose Ectrepopterus (ordered according to their consistency indexes). Characters 62, 67, 168, and 310 were also found as unambiguous autapomorphies of Ectrepopterus in the implied weighting analysis. - (Ch. 168: 2; ci = 0.33) Foramen in posterior region of metapterygoid that serves as passage for the ramus mandibularis of the trigeminus nerve forms an incomplete arch, bordered posteriorly by the hyomandibula (Fig. 2). This foramen is situated entirely within the metapterygoid among characid fishes lacking a supraorbital. The condition found in Ectrepopterus is apomorphic, and a reversal, since the foramen opening posteriorly is found in several basal lineages of the Characidae (e. g. Brycon M��ller and Troschel 1844, Bryconops Kner 1858, and Iguanodectes Cope 1872; see Mirande 2010). This condition was found homoplastically in Bryconamericus scleroparius (Regan 1908) and in Pseudocorynopoma doriae Perugia 1891. - (Ch. 310: 1; ci = 0.10) Pectoral-fin rays bearing hooks. Although the presence of hooks in the fin rays is a condition shared by most characid fishes (Malabarba and Weitzman 2003), its presence in the pectoral fin rays is not usual and was found by parsimony as diagnostic for Ectrepopterus. It was found homoplastic in Astyanax cf. asuncionensis G��ry 1972, A. chico Casciotta and Almir��n 2004, A. lineatus (Perugia 1891), A. troya Azpelicueta, Casciotta and Almir��n 2002, Bario steindachneri (Eigenmann 1893), Bryconamericus iheringii (Boulenger 1887), B. rubropictus (Berg 1901), B. thomasi Fowler 1940, Hyphessobrycon luetkenii (Boulenger 1887), H. socolofi Weitzman 1977, and Nematocharax venustus Weitzman, Menezes and Britski 1986 (Mirande 2010). - (Ch. 62: 2; ci = 0.06) Posterior margin of second infraorbital posteroventrally oblique and second infraorbital ventrally bordering anterior region of third infraorbital (Fig. 3). This condition is usually associated with a long maxilla, and is found homoplastic in Exodon paradoxus M��ller and Troschel 1844, Hollandichthys multifasciatus (Eigenmann and Norris 1900), Oligosarcus spp., and several genera of the Characinae. - (Ch. 67:0; ci = 0.03) Fourth infraorbital (Fig. 3) more developed longitudinally than dorsoventrally (versus longer dorsoventrally than longitudinally). Although apomorphic for Ectrepopterus, this character is broadly distributed among Characidae. - (Ch. 104: 1; ci = 0.03) Ascending process of premaxilla reaching just anterior end of nasal (versus reaching at least one-third of length of nasal). Although apomorphic for Ectrepopterus, this character is broadly distributed among Characidae. - (Ch. 91: 1; ci = 0.02) Lateral line interrupted (versus complete). Although highly variable in Characidae, this character was recovered as one of the autapomorphies for Ectrepopterus., Published as part of Malabarba, Luiz R., Bertaco, Vinicius A., Carvalho, Fernando R. & Litz, Thomas O., 2012, Revalidation of the genus Ectrepopterus Fowler (Teleostei: Characiformes), with the redescription of its type species, E. uruguayensis, pp. 47-60 in Zootaxa 3204 on pages 49-50, DOI: 10.5281/zenodo.210066, {"references":["Fowler, H. W. (1943) Notes and descriptions of new or little known fishes from Uruguay. Proceedings of the Academy of Natural Science of Philadelphia, 95, 311 - 334.","Gery, J. (1972) Corrected and supplemented descriptions of certain characoid fishes described by Henry W. Fowler, with revisions of several of their genera. Studies on the Neotropical Fauna, 7, 1 - 35.","Gery, J. (1977) Characoids of the World. T. H. F. Publications, Neptune City, NJ, 672 p.","Weitzman, S. H. & Palmer, L. (1997) A new species Hyphessobrycon (Teleostei: Characidae) from the Neblina region of Venezuela and Brazil, with comments on the putative ' rosy tetra clade'. Ichthyological Exploration of Freshwaters, 7, 209 - 242.","Mirande, J. M. (2010) Phylogeny of the family Characidae (Teleostei: Characiformes): from characters to taxonomy. Neotropical Ichthyology, 8, 385 - 568.","Malabarba, L. R. & Weitzman, S. H. (2003) Description of new genus with six new species from southern Brazil, Uruguay and Argentina, with a discussion of a putative characid clade (Teleostei: Characiformes: Characidae). Comunicacoes do Museu de Ciencias e Tecnologia da PUCRS, Serie Zoologia, 16, 67 - 151."]}

Moenkhausia aurantia, new species (Figs. 1���3; Table 1) Holotype. UFRGS 13619, 40.6 mm SL, female, Brazil, Goi��s, Nova Roma, c��rrego Kavanca at fazenda Kavanca, upper rio Tocantins basin, 13 �� 41 ��� 34 ������S 46 �� 57 ��� 45 ������W, V. A. Bertaco, F. R. Carvalho & G. L. C. Frainer, 9 Sep 2009. Paratypes. Brazil: Goi��s: Nova Roma, upper rio Tocantins basin: UFRGS 11224, 15 (9), 24.2���37.5 mm SL, UNT 10052, 8 (3), 27.5���36.4 mm SL, stream on GO 112 at locality Lagoa Brava, between Nova Roma and Teresina de Goi��s, rio Paran�� basin, 13 �� 48 ��� 37 ������S 46 �� 51 ��� 57 ������W, V. A. Bertaco, F. R. Carvalho & G. L. C. Frainer, 9 Sep 2009. UFRGS 11251, 23 (7), 5 c&s, 15.6���45.5 mm SL, INPA 34990, 8 (3), 29.2���37.8 mm SL, MCP 45865, 8 (4), 24.3���38.4 mm SL, MZUSP 107827, 8 (3), 27.7 ���40.0 mm SL, c��rrego Kavanca at fazenda Kavanca, 13 �� 41 ��� 34 ������S 46 �� 57 ��� 45 ������W, V. A. Bertaco, F. R. Carvalho & G. L. C. Frainer, 9 Sep 2009. Diagnosis. Moenkhausia aurantia can be distinguished from all its congeners by the combination of 3 or 4 scale rows below and 5 or 6 scale rows above the lateral line, 21���25 branched anal-fin rays, 33���35 lateral line scales, 2���4 maxillary teeth, 9���12 dentary teeth, and color pattern composed by hyaline caudal-fin lobes, one diffuse humeral spot, and a one-scale-depth midlateral stripe continuous to the caudal peduncle spot. Among Moenkhausia species, M. aurantia is similar to M. lopesi, from rio Cuiab�� basin in the upper rio Paraguay system. It differs from M. lopesi by the lower number of maxillary teeth (2���4 vs. 3���7), total number of dentary teeth (9���12 vs. 14���19), body depth (34.2���40.6 vs. 30.2 ���34.0% of SL), anal-fin base length (30.8���35.7 vs. 27.4���31.7 % of SL), distal profile of the anal fin (straight to convex vs. deeply concave), and the presence of a faint humeral spot (vs. absent). Furthermore, M. aurantia is distinguished from all congeners in the rio Tocantins-Araguaia basin by the number of lateral line scales (33���35 vs. 27���28 in M. pyrophthalma Costa, 36���37 in M. dasalmas Bertaco, Jerep & Carvalho, 36���38 in M. loweae G��ry, and 37���41 in M. pankilopteryx Bertaco & Lucinda), by the number of branched anal-fin rays (21���25 vs. 17���19 in M. pyrophthalma and M. dasalmas, 26���30 in M. tergimacula Lucena & Lucena), by the number of humeral spots (1 vs. 2 in M. pankilopteryx and M. dasalmas), by the absence of a black spot in the upper caudal-fin lobe (vs. present in M. hysterosticta Lucinda, Malabarba & Benine and M. loweae), and by the absence of a dark spot, saddle-like mark anterior to the dorsal-fin origin (vs. present in M. tergimacula). See Discussion for detailed comparison with the remaining species of Moenkhausia. Description. Morphometric data are summarized in Table 1. Body compressed, moderately short, greatest body depth usually located anterior to dorsal-fin origin. Dorsal profile of head convex from tip of upper jaw to vertical through anterior nostril; slightly straight or convex from that point to tip of supraoccipital spine. Dorsal profile of body slightly convex from posterior tip of supraoccipital spine to base of last dorsal-fin ray, and straight to adipose-fin origin. Ventral profile of body convex from tip of lower jaw to pelvic-fin origin, straight or slightly convex from that point to anal-fin origin, and straight and posterodorsally slanted along anal-fin base. Dorsal and ventral profile of caudal peduncle straight to slightly concave. The range includes the holotype. SD = Standard deviation. Mouth terminal, jaw isognathous. Maxilla extending posteroventrally to vertical through anterior half of orbit, aligned approximately at 45 degree angle relative to longitudinal axis of body. Main axis of maxilla straight, with approximately same width along all its length. Two tooth rows in premaxilla: outer row with three* or four tri- to pentacuspid teeth (mode = 4, n = 30), central cusp longer; inner row with five teeth, gradually decreasing in length from first to fourth, last distinctly smaller, with five cusps; central cusp twice as long and broad as others cusps. Maxilla with two to four teeth (3 *, mode = 3, n = 30), three to five cusps, with central cusp slightly longer. Four anteriormost dentary teeth larger, with five cusps, followed by five or eight teeth with one to three cusps. Central cusp in all teeth twice or three times longer and broader than lateral cusps. All cusp tips slightly curved posteriorly towards oral cavity (Fig. 2). Dorsal-fin rays ii, 9 * (n = 30); first unbranched ray approximately half-length of second ray. First and second branched rays longest. Dorsal-fin origin posterior to middle of body, at vertical through anterior third of pelvic-fin base. Tip of dorsal fin reaching adipose-fin origin in specimens larger than 35.0 mm SL. Distal margin of dorsal fin nearly straight to somewhat convex. Adipose fin origin at vertical through fourth or fifth last anal-fin rays insertion. Anal-fin rays iv���v, 21���25 (iv, 24 *, mode = iv, 23, n = 30). First unbranched ray usually visible only in c&s specimens. Distal profile of anal fin distal profile smoothly concave. Anal fin origin at vertical through base of two or three last dorsal-fin rays. Pectoral-fin rays i, 11���12 * (mode = 11, n = 30). Tip of pectoral fin surpassing pelvic-fin origin. Pelvic-fin rays i, 7 * (n = 30). Pelvic-fin origin slightly anterior to vertical through dorsal-fin origin. Tip of pelvic fin reaching and surpassing anal-fin origin. Caudal fin forked, lobes similar in size, 19 * principal rays (n = 30). Dorsal and ventral procurrent caudal-fin rays 9���11 and 8���10, respectively (n = 5). Lateral line complete, with 33���35 * perforated scales (34 *, mode = 34, n = 30). Scale rows between dorsal-fin origin and lateral line 5 or 6 * (mode = 6, n = 30); scale rows between lateral line and pelvic-fin origin 3 or 4 * (mode = 4, n = 30). Predorsal scales 10 *��� 12 (mode = 11, n = 30) arranged in regular series. Scale rows around caudal peduncle 14 * (mode = 14, n = 30). Axillary scale on pelvic fin origin covering 1 or 2 scales posteriorly. Scale sheath along anal-fin base 8���10 * scales (mode = 10, n = 30), in single series, covering base of anteriormost rays. Caudal fin scaled, scales over first third of upper and along half-length of lower caudal-fin lobes; scales gradually decreasing in size posteriorly. Precaudal vertebrae 15 or 16; caudal vertebrae 17���19; total vertebrae 33 or 34 (n = 5). Supraneurals 4 (n = 5), distal portion widened. Branchiostegal rays 4 (n = 5). First gill arch with six rakers on epibranchial, one between epibranchial and ceratobranchial, eight to nine on ceratobranchial, and two on hypobranchial (n = 5). Color in alcohol. Overall body color varying from light to pale yellowish. Head dusk to dark brownish dorsally. Region from the second to fifth infraorbital and opercular apparatus silvery. Several small dark melanophores surrounding nares, lips, maxillary, lower jaw and anterior margin of orbit, becoming sparse posteriorly; larger melanophores scattered over surface of infraorbitals 3���6 and opercular series on larger specimens. Overall iris color silvery, darker by presence of black pigment bellow ventral margin of pupil and at dorsal portion of eye. High concentration of melanophores in humeral region, forming inconspicuous vertically elongated humeral spot. Dorsum line darker than body lateral and ventral regions. Dark melanophores scattered over body, except ventral region from lower jaw articulation to urogenital papillae. Dorsal and dorso-lateral scales exhibiting tenuous reticulated pattern due to higher concentration of melanophores at their distal borders. Melanophores distributed along dark longitudinal line, larger than other body melanophores. Subcutaneous longitudinal dark stripe starting thinner at vertical line just anterior to dorsal-fin origin, widening posteriorly up to base of caudal fin, reaching one scale wide. Caudal peduncle with higher concentration of melanophores shaping a caudal black spot continuous to longitudinal line, and extending posteriorly over middle caudal-fin rays. Few melanophores following myoseptum of muscles hypaxialis at lower lateral side of caudal region. Dorsal, pectoral and pelvic fins hyaline with scattered melanophores at interradial membrane and bordering margins of rays. Anal and caudal also hyaline, with scattered melanophores more concentrated at distal margins of those fins (Fig. 1). Color in life. Overall body and head color pattern silvery to whitish. Melanophore distribution pattern of head, body and fins as described above for alcohol preserved specimens. Body slightly orangish, with higher intensity in ventral regions of head, abdomen, base of anal fin and caudal peduncle. Females with less intense orangish color. Dorsal fin orangish, more intense pigmented at first half length of rays. Distal half of second unbranched and first branched dorsal-fin rays hyaline to whitish. Pectoral fin hyaline. Pelvic fin orangish at its proximal portion. Anal fin orangish, more intense along proximal half of the most anterior anal-fin rays. Distal third of last unbranched and first two branched anal-fin rays hyaline to whitish. Adipose fin orangish, more intense colored at its proximal portion. Caudal fin presenting orange coloration at proximal half of dorsal and ventral lobes rays, middle caudal-fin rays dark by presence of melanophores (Fig. 3). Sexual dimorphism. Secondary sexual characters were not found on examined specimens. Mature gonads were observed in five dissected and c&s specimens (UFRGS 11251, two males, 35.4 and 36.0 mm SL; three females, 33.2���36.7 mm SL). Distribution. Moenkhausia aurantia is known from tributaries of the rio Paran��, upper rio Tocantins basin, in the Chapada dos Veadeiros region, Brazilian Cerrado, Goi��s, Brazil (Fig. 4). Etymology. The specific epithet is derived from the Latin aurantium, employed here as an adjective in reference to the distinctive orangish coloration of this species. Ecological notes. The type locality is around 600 m above sea level. Moenkhausia aurantia inhabits streams, and occurs in semi-lentic and lotic shallow areas (up to 0.8 m deep) with riparian vegetation composed by trees and shrubs (Fig. 5). The streams have transparent water, and bottom with rocks, stones, and sand in some stretches. The new species was syntopically collected with Ancistrus sp., Aspidoras albater, Astyanax sp., A. elachylepis, Characidium sp., Cichlasoma araguaiense, Creagrutus britskii, Eigenmannia sp., Hasemania sp., Harttia sp., Hemiodus cf. ternetzi, Hypostomus sp., Imparfinis sp., Ituglanis sp., Knodus sp., Leporinus sp., Moenkhausia sp., Pimelodella sp., and Steindachnerina amazonica. Stomach contents of five specimens (UFRGS 11251) were mainly composed by Hymenoptera, Hirudinea, filamentous alga and digested vegetal organic matter., Published as part of Bertaco, Vinicius A., Jerep, Fernando C. & Carvalho, Fernando R., 2011, A new characid fish, Moenkhausia aurantia (Ostariophysi: Characiformes: Characidae), from the upper rio Tocantins basin in Central Brazil, pp. 29-38 in Zootaxa 2934 on pages 30-34, DOI: 10.5281/zenodo.278082

Key to the species of Moenkhausia described from Tocantins-Araguaia basin 1 Branched anal-fin rays 17 to 19.......................................................................... 2 - Branched anal-fin rays 21 to 30.......................................................................... 3 2 Lateral line scales 27 or 28; one maxillary tooth; presence of a dark blotch on caudal-fin base; black triangular humeral spot......................................................................................... M. pyrophthalma - Lateral line scales 36 or 37; four to five maxillary tooth; absence of a dark blotch on caudal-fin base; humeral spot diffuse.............................................................................................. M. dasalmas 3 Dark spot on upper caudal-fin lobe....................................................................... 4 - Absence of dark spot on upper caudal-fin lobe............................................................... 5 4 Lateral line scales 29 to 35; a large and irregularly shaped humeral spot located above the fifth to ninth perforated scale of lateral line............................................................................... M. hysterosticta - Lateral line scales 36 to 38; a vertically elongate humeral spot located just after upper margin of opercle......... M. loweae 5 Two humeral spots; 7 or 8 scales above and 6 or 7 scales below the lateral line; 37 to 41 lateral line scales.. M. pankilopteryx - One humeral spot; 5 to 6 scales above and 3 or 4 scales below the lateral line; 33 to 37 lateral line scales............... 6 6 Branched anal-fin rays 21 to 25; predorsal scales 7 or 8; presence of a dark saddle-like mark anterior to the dorsal-fin origin; lateral line scales 35 to 37.................................................................. M. tergimacula - Branched anal-fin rays 26 to 30; predorsal scales 10 to 12; absence of dark spot anterior to dorsal-fin origin; lateral line scales 33 to 35......................................................................... M. aurantia, new species, Published as part of Bertaco, Vinicius A., Jerep, Fernando C. & Carvalho, Fernando R., 2011, A new characid fish, Moenkhausia aurantia (Ostariophysi: Characiformes: Characidae), from the upper rio Tocantins basin in Central Brazil, pp. 29-38 in Zootaxa 2934 on page 37, DOI: 10.5281/zenodo.278082

Hemigrammus parana, new species (Fig. 1) Holotype. DZSJRP 6217, 24.9 mm SL, rio Grande near the old sand port, known as Velho Ad��o, municipality of Santa Clara D���Oeste, S��o Paulo state, Brazil, 20 ��01��� 27.2 ���S, 50 �� 54 ��� 46.6 ���W, F.R.Carvalho and J. O.Carvalho, 08.i. 2004. Paratypes. All from Brazil, upper rio Paran�� system. DZSJRP 8781, 1262/ 3, 13.3 ���28.0 mm, Ribeir��o Can-Can, old port, Ilha Solteira reservoir, municipality of Santa Clara D��Oeste, S��o Paulo state, 20 ��01��� 26.5 ���S 50 �� 54 ��� 44.5 ���W, F.Langeani, R.S.Costa-Ferreira, F.R.Carvalho, M.M.F.Marinho, J.P.Serra and F.L.R.Sousa, 23.viii. 2006; DZSJRP 8809, 208 / 5, 12.2���30.8 mm, headwater of Brejo Comprido, Ilha Solteira reservoir, municipality of Aparecida do Taboado, Mato Grosso do Sul state, 20 �� 10 ��� 34.4 ���S 51 �� 12 ��� 18.6 ���W, F.Langeani, R.S.Costa-Ferreira, F.R.Carvalho, M.M.F.Marinho, J.P.Serra and F.L.R.Sousa, 26.viii. 2006; DZSJRP 9051, 1399, 11.8���29.5 mm, C��rrego da Anta, Fazenda S��o F��lix, Ilha Solteira reservoir, municipality of Aparecida do Taboado, Mato Grosso do Sul state, 20 �� 23 ��� 57.4 ���S 51 ��07��� 57.3 ���W, F.Langeani, R.S.Costa-Ferreira, F.R.Carvalho, M.M.F.Marinho, J.P.Serra and F.L.R.Sousa, 29.viii. 2006; DZSJRP 10796, 359 / 34, 14.8���26.1 mm, 3 c&s, same data as holotype; LIRP 6004 (ex- DZSJRP 10796), 20, 19.8 ���25.0 mm; MCP 41759 (ex- DZSJRP 8781), 20, 20.7���25.2 mm; MNRJ 31197 (ex-DZSJRP 8781), 20, 20.7���27.9 mm; MZUEL 4904 (ex- DZSJRP 10796), 20, 25.7 ��� 18.8 mm; MZUSP 95001 (ex-DZSJRP 8781), 30, 19.3���26.8 mm; UFRGS 8869 (ex-DZSJRP 9051), 50 / 16, 22.2���26.7 mm. Diagnosis. Hemigrammus parana differs from all congeners, except Hemigrammus levis, by the absence of a humeral spot and presence of a conspicuous black spot, restricted to caudal fin, roughly triangular or rectangular, extending from base to tip of middle caudal-fin rays (its greatest depth at base of caudal-fin rays). It differs from H. levis by having 18���23 (mode 21) unbranched anal-fin rays (vs. 16���18), largest tooth of the inner row of premaxilla and dentary with 5 cusps (vs. 7���9 cusps), anterior portion of longitudinal band as a broad uninterrupted pigmented area (vs. anterior portion of longitudinal band with a small round concentration of chromatophores at humeral region, surrounded by unpigmented areas anterior and posteriorly), distance of snout to anal-fin origin 58.6���65.9 % SL, mean 62.0% (vs. 63.7���73.9 % SL, mean 68.8 %), anal base length 25.1���32.5 % SL, mean 29.2 % (19.3���24.7 % SL, mean 22.0%), upper jaw length 39.6���48.3 % HL (vs. 34.3���39.5 % HL), Description. Morphometric data presented in Table 1. Overall size small (22.2���30.2 mm SL). Body compressed, moderately elongate, greatest body depth at vertical through dorsal-fin origin. Dorsal profile of head convex from tip of upper jaw to vertical through anterior nostril; straight to slightly concave from that point to tip of supraoccipital spine. Dorsal profile of body slightly convex from posterior tip of supraoccipital spine to dorsal-fin origin; straight and posteroventrally inclined from dorsal-fin origin to adipose-fin origin and slightly concave along caudal peduncle. Ventral profile of body convex from tip of lower jaw to pelvic-fin origin; straight from that point to anal-fin origin; straight and posterodorsally slanted along anal-fin base. Ventral profile of caudal peduncle slightly concave. Eyes relatively large, lacking distinct adipose eyelid. Jaw equal, mouth terminal. Maxillary extending posteriorly anterior margin of eye, slightly curved, aligned approximately at 45 degrees angles relative to longitudinal axis of body. Premaxillary teeth in two rows: outer with 2 (7), 3 (29), 4 *(20), or 5 (3) tricuspid teeth (rarely pentacuspid); inner tooth row with 5 tri-pentacuspid teeth. Dentary with 3 (2), 4 *(54) or 5 (2) large tri to pentacuspid teeth, followed by a series of 5���10 small conical teeth. Antero and posterodorsal border of maxillary relatively straight. Maxilla with 0(5), 1 *(29), 2 (17), or 3 (5) tricuspid teeth along anteroventral margin (Fig. 2). Central median cusp in all teeth longer than remaining cusps; cusp tips slightly curved posteriorly and towards inside of mouth. Scales cycloid, with few radii, relatively small, and weakly implanted. Lateral line incomplete, slightly decurved anteriorly to vertical through distal tip of pectoral fin; perforated scales of lateral line 6 (7), 7 (19), 8 *(15), 9 (13), 10 (2), or 11 (1); longitudinal scales series including pored scales 26 (2), 27 (4), 28 (6), 29 (8), 30 (6), 31 (10), 32 *(12), 33 (6), or 34 (1); 5 (58) scales rows between dorsal-fin origin and lateral line; scales rows between lateral line and pelvic-fin insertion 3 (18) or 4 *(32). Predorsal scales 9 (13), 10 *(40), or 11 (6) scales. Scale sheath along anal-fin base with 2 (2), 3 (10), 4 (20), 5 (14), or 6 *(1) in a single row. Circumpeduncular scales 10 *(14), 11 (35) or 12 (6). Axillary scale present. Dorsal-fin rays ii, 9 (58); first unbranched ray approximately one-half of second unbranched ray. Dorsalfin origin at midbody, at vertical through anterior third of pelvic- fin base, and base of its last ray at vertical through second or third anal-fin branched ray. Tip of longest ray of adpressed dorsal fin at vertical through 11 th and 12 th branched anal-fin rays. First dorsal-fin pterygiphore inserting behind neural spine of 6 th(2) or 7 th(1) vertebra. Adipose fin present, located approximately at vertical through second or third last branched of analfin rays. Pectoral fin with i, 9 (2), 10 (17), 11 *(37), or 12 (3) rays. Pelvic fin with i, 6 (7) or 7 *(52) rays; tip of longest ray reaching anal fin. Anal-fin rays iii, 18 (1), 19 (9), 20 (14), 21 *(17), 22 (14), or 23 (4). Caudal fin forked, lobes slightly rounded, similar in size, with i, 9, 8, i rays (58). Caudal fin partially covered with very small and weakly implanted scales along first third of upper and first half of lower caudal-fin lobes. Dorsal procurrent caudal-fin rays 10 (3), ventral procurrent caudal-fin rays 8 (2) or 9 (1). Branchiostegal rays 4. First gill arch with 6 gill rakers on epibranchial, 10 in ceratobranchial, and 3 on hypobranchial. Precaudal vertebrae 15 and caudal vertebrae 17 (2) or 18 (1). Supraneurals 4. Color in alcohol. Overall ground coloration of body pale. Infraorbital and opercular areas silvery. Anterior portion of lower jaw, dorsal portion head, snout, and dorsal midline of body covered by small dark chromatophores. Longitudinal body band dusky, originating as a broad pigmented area from opercle to vertical through tip of longest pectoral-fin ray, becoming more intense, slender and sometimes silvery from this point to end of caudal peduncle. Scales of second or third longitudinal rows above lateral line bordered by few chromatophores, forming a slight reticulate pigmentation. Humeral spot absent. Dorsal, adipose, pectoral, pelvic, and anal fins almost entirely hyaline, with few scattered dark cromatophores. Posterior half of caudal-fin lobes with diffuse chromatophores, making fin slightly darkish. Conspicuous black caudal spot roughly triangular or rectangular, extending from base to tip of middle caudal-fin rays, its greatest depth at base of caudal rays; sometimes, caudal spot lighter, not reaching the tip of middle caudal-fin rays. Color in life. Overall coloration silvery. Intense silvery longitudinal band on body. Dorsal, adipose, pectoral, pelvic, and anal fins yellowish; median portion of caudal lobes reddish or orange. Conspicuous black caudal spot, extending from base to tip of middle caudal-fin rays. Sexual dimorphism. Secondary sexually dimorphic characters, such as hooks on pelvic and anal-fin rays, were not found on examined specimens. Distribution. Hemigrammus parana only occurs in the area of influence of the Ilha Solteira reservoir, upper Paran�� system, southeastern Brazil, throughout margins of rio Grande, rio Paran��, rio Parana��ba, and rio S��o Jos�� dos Dourados (Fig. 3 ��� 4). Etymology. The specific name parana refers to the type locality river, the rio Paran��, which originates after the confluence of the rio Grande and the rio Parana��ba, where the new species occurs. Ecological notes. Hemigrammus parana is very common and specially abundant in backwater margins, from 0.3 to 1.30 m depth, occurring associated with Poaceae and macrophytes (Ceratophyllum sp., Egeria densa, Eichornia spp., Ludwigia sedorde, and Salvinia sp.). Stomach contents examination of two specimens showed a large volume of Spirogyra sp., Cladocera, sediments (sand) and unidentified organic matter. Several species were collected syntopically with Hemigrammus parana, the most frequent were Astyanax altiparanae Garutti & Britski, Crenicichla britskii Kullander, Geophagus proximus (Castelnau, 1855), Gymnotus carapo Linnaeus, 1758, Hoplias malabaricus (Bloch), Hyphessobrycon eques (Steindachner), Laetacara sp., Metynnis maculatus (Kner), Moenkhausia sp., Pamphorichthys hollandi (Henn), Roeboides descalvadensis Fowler, Satanoperca pappaterra (Heckel), Serrapinnus notomelas (Eigenmann), and Serrasalmus maculatus Kner., Published as part of Marinho, Manoela M. F., Carvalho, Fernando R., Langeani, Francisco & Tatsumi, Vio L., 2008, A new Hemigrammus Gill from upper rio Paran�� system, Southeastern Brazil (Characiformes: Characidae), pp. 52-60 in Zootaxa 1724 on pages 53-56, DOI: 10.5281/zenodo.181167

Made available in DSpace on 2022-04-28T20:07:38Z (GMT). No. of bitstreams: 0 Previous issue date: 2006-03-01 We investigated sex ratio, reproductive period, fecundity, and sexual dimorphism of Pamphorichthys hollandi in southeastern Brazil. Reproductive status was identified according to a female's gonads/embryos development scale. Of the 613 collected specimens, 373 (60%) were females and 240 (40%) males giving a sex ratio of 1 male:1.6 female. Sexual dimorphism was evident in respect to body size, body coloration, and anal and pelvic-fin morphologies. The highest proportion of reproductive females was registered in January and February, which represent wet and hot periods. The simultaneous presence of more than one litter at different stages of development within a single female characterized this species as superfetaceous. © 2006 by Verlag Dr. Friedrich Pfeil. UNESP Universidade Estadual Paulista Departamento de Zoologia e Botânica, Rua Cristovão Colombo, 2265, 15054-000, Sao Jose do Rio Preto, SP UNESP Universidade Estadual Paulista Departamento de Zoologia e Botânica, Rua Cristovão Colombo, 2265, 15054-000, Sao Jose do Rio Preto, SP