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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