Gephyrocharax melanocheir Eigenmann, 1912

Vanegas-Ríos, James A., 2016, Taxonomic review of the Neotropical genus Gephyrocharax Eigenmann, 1912 (Characiformes, Characidae, Stevardiinae), Zootaxa 4100 (1) : 52-58

Gephyrocharax melanocheir Eigenmann, 1912

( Figs. 22–24 )

Gephyrocharax melanocheir Eigenmann, 1912: 24 –25 [original description, holotype FMNH 56049 (CM 4840, type lost), type locality: Bernal creek (= Colombia: Tolima, Bernal creek near Honda)]. Eigenmann, 1914: 41 [listed, key]. Eigenmann, 1920c: 32 [listed from Magdalena River basin]. Myers in Eigenmann & Myers, 1929: 477, 480–481, pl. 63, fig. 3 [redescribed without figure]. Schultz, 1944: 322 [key]. Fowler, 1943: 243 [ Colombia: Magdalena department, acequia in Aracataca]. Fowler, 1945: 5 [ Colombia: Honda, Magdalena River basin]. Miles, 1947: 156, fig. 106 [listed from Magdalena River, key]. Bussing & Roberts, 1971: 179 –180 [comments on morphology of males]. Dahl, 1971: 133 –134 [key, distributional data]. Bussing, 1974: 136 [comments on morphology of males]. Mojica, 1999: 557 [listed from Colombia, data compilation]. Bushmann, Burns & Weitzman, 2002: 189 [gill gland morphology]. Weitzman, 2003: 225 [catalogue]. Burns & Weitzman, 2005: 122, 127 [comments on pectoral fin of males, evidence of insemination]. Maldonado-Ocampo, Ortega-Lara, Usma, Galvis, Villa-Navarro, Vásquez, Prada-Pedreros & Ardila, 2005: 90, fig. 93 [recorded for Andean drainages from Colombia, ecological and morphological data]. Mojica, Castellanos, Sánchez-Duarte & Díaz, 2006a: 135 [listed from Ranchería River]. Mojica, Galvis, Sánchez-Duarte, Castellanos & Villa-Navarro, 2006b: [listed from middle Valley of Magdalena]. Villa-Navarro, Zuñiga-Upegui, Castro-Roa, García-Melo, García-Melo & Herrada-Yara, 2006: 10: [listed from upper Magdalena River]. Maldonado-Ocampo, Vari & Usma, 2008: 181 [listed from Colombia]. Álvarez-León, Orozco-Rey, Páramo-Fonseca & Restrepo-Santamaría, 2013: 101 [listed from Colombia]. Bonilla-Rivero & López-Rojas, 2013: 489, fig. 1 [distribution map]. Vanegas-Ríos, Azpelicueta, Mirande, Gonzales, 2013: 282 –283, fig. 7 [compared with G. t o r re s i, examined material]. Thomaz, Arcila, Ortí & Malabarba, 2015: Add. File 5 [tentative classification].

Gephyrocharax chocoensis [not of Eigenmann, 1912]. Fowler, 1945: 6 [misidentified].

Diagnosis. Gephyrocharax melanocheir differs from its congeners by having an intense dark pigmentation at the base of the five anterior dorsal-fin rays (vs. absence of this pigmentation) and the fan-shaped structure distally formed by the highly developed branching pattern of the outermost branched pectoral-fin ray of adult males (vs. this pectoral-fin ray with less extensive branching pattern, similar to that of contiguous rays, not distally forming such structure, except in G. t o r re s i). This fan-shaped structure has minute bony hooks and presents distally a dark blotch (some specimens with a whitish tonality); both features are absent in the remaining congeners, except in G. martae , which has the distal portion of the pectoral-fin ray similarly pigmented. The lack of terminal lateral-line tube between caudal-fin rays 10 and 11 (vs. possession of this tube) differentiates the species from almost all its congeners, except G. sinuensis (absent in two specimens), G. t o r res i, and G. valencia . Gephyrocharax melanocheir is also distinguished from G. atracaudatus , G. chocoensis , G. intermedius , G. major , and G. venezuelae by the gillgland length of adult males (7.0–11.4 % SL vs. 0.5–5.4 % SL) and the externally developed urogenital papillae in adult females (vs. this papillae not developed externally). From G. caucanus , G. chocoensis , G. sinuensis , G. t o r re s i, and G. venezuelae , G. melanocheir differs by lacking osseous lamella between the first and second basibranchials (vs. possession of this lamella). Gephyrocharax melanocheir is further distinguished from G. torresi by the snout length (22.1–28.4 % HL vs. 28.3–31.8 % HL), number of vertebrae (38–39 vs. 40–41), number of terminal minute branches forming the fan-shaped structure of pectoral fin in adult males (28–54 vs. 60–88), the posterior margin of mesethmoid concave on its central region (vs. straight), the pouch scale of adult males dorsally reaching to caudalfin ray 10 or an area located between caudal-fin rays 9 and 10 (vs. dorsally reaching to ray 11 or an area located between rays 11 and 12), and the three or rarely four interradialis bundles on lower caudal-fin lobe posteriorly inserting beyond the posterior margin of the pouch scale in adult males (vs. one or two).

Description. Morphometric data in Table 6 . Largest male 39.3 mm SL, largest female 41.9 mm SL. Body laterally compressed, with maximum depth at vertical point between anal- and pelvic-fin origins. Dorsal profile of head slightly convex from margin of upper lip to tip of supraoccipital spine. Dorsal profile of body straight or slightly convex from that point to dorsal-fin origin, slanting posteroventrally along dorsal-fin base, and straight from posteriormost dorsal-fin ray to caudal peduncle ( Fig. 22 ). Ventral profile of body convex from tip of dentary to pelvic-fin origin, straight from that point to anal-fin origin, and slanting posterodorsally and straight or slightly convex from this point to caudal peduncle origin. Anterior fontanel present, usually well developed, rarely reduced to narrow opening due to anterior contact between frontals. Anterior nostril rounded and separated by skin fold from larger posterior nostril. Groove with at least three rows of neuromasts extending from half-length between posterior pore of nasal bone and nostrils to posterior portion of frontals. Small groove with few neuromasts between nostrils and nasal bones.

Mouth superior, lower jaw projecting anterior to tip of upper jaw. Premaxilla with two rows of teeth. Outer row with 2 (5), 3 (19), 4* (58), or 5 (2) teeth; usually tricuspid, rarely bi- or tetracuspid teeth. Inner row with 4 (24), 5* (59), or 6 (1) teeth; symphyseal tooth tri- or tetracuspid and remaining teeth penta- or hexacuspid. Maxilla sometimes toothless (2), usually with 1* (74), 2 (6), or 3 (2) teeth; bi- or pentacuspid, dorsalmost tooth often with greater number of cusps ( Fig. 23 A). Maxilla posteriorly reaching vertical through anterior one-third of eye. Dentary with 7 (1), 8 (1), 9* (8), 10 (3), 11 (11), 12 (17), 13 (15), 14 (7), or 15 (1) teeth; three anteriormost teeth larger, pentacuspid, followed by one median-sized tooth tri- to pentacuspid, and 3, (1), 4 (1), 5* (8), 6 (3), 7 (3), 8 (11), 9 (17), 10 (7), or 11 (1) smaller conical to tricuspid teeth ( Fig. 23 B).

Dorsal-fin rays ii,7 (4), 8* (83), or 9 (1). Nine* proximal pterygiophores on dorsal fin (8 rad, 6 c&s). Dorsalfin origin located at vertical between branched anal-fin rays 4–9. Adipose-fin origin located at vertical through bases of two posteriormost anal-fin rays. Anal-fin rays iii (1), iv* (54), v (37), or vi (1), 24 (2), 25 (12), 26* (24), 27 (29), 28 (14), or 29 (12) ( Fig. 6 ). Twenty-six to 30 proximal pterygiophores on anal fin (7 rad, 6 c&s). Anal-fin origin closer to origin of hypural joint than to snout tip. Pectoral-fin i,8 (1), 9 (50), 10* (33), or 11 (6). Pectoral-fin distal tip posteriorly reaching one-fifth to pelvic-fin distal tip. Pelvic-fin rays i,6* (79) or 7 (4) (one atypical specimen with 5 branched rays). Pelvic-fin origin at vertical between pored lateral-line scales 8–9 and slightly anterior to body midlength. Caudal fin forked with 10/9 principal rays in all specimens.

Scales cycloid, with several radii along posterior margin. Lateral line complete, pored scales 37 (7), 38* (19), 39 (32), 40 (22), 41 (8), 42 (5), or 43 (1). Terminal lateral-line tube absent. Predorsal scales 17 (15), 18* (53), 19 (18), or 20 (4). Scale rows between dorsal fin and lateral line 5* (33), 6 (57), or 7 (1). Scale rows between lateral line and anal fin 4 (16), 5* (74), or 6 (1). Scale rows between lateral line and pelvic fin 3 (2), 4* (45), or 5 (39). Circumpeduncular scales 14* (54), 15 (30), or 16 (2). One row of scales forming sheath along anal-fin base; main ventral row with 11 (6), 12 (7), 13 (6), 14 (8), 15 (13), 16 (14), 17 (6), 18 (7), 19 (1), 20 (1), 21 (1), or 23 (2) scales. Total number of vertebrae 38* (5) or 39 (8); 15* (11) precaudal and 23* (4) or 24 (7) caudal (8 rad, 6 c&s). Gillrakers on dorsal limb of first branchial arch 5(4), 6 (37), or 7 (11); ventral limb with 11 (15), 12 (35), or 13* (4) (upper gill rakers not counted in neotype).

Color in alcohol. Ground color pale yellowish or light brown, darker along mid-dorsal line and slightly lighter ventrally. Minute dark chromatophores scattered on body, less numerous on lateral region of abdomen. Dark midlateral stripe diffuse (silvery in specimens retaining guanine: CZUT-IC 7883), extending from vertical through pelvic-fin origin to middle or posterior region of caudal peduncle. Dark chromatophores present along myosepta between lateral line and upper portion of anal fin. Dark humeral blotch vertically elongate and narrow. Dark blotch on caudal peduncle, horizontally elongate, and extending from middle region of caudal peduncle to caudal-fin rays. Dorsal fin light gray, with few dark chromatophores on rays and membranes. Bases of five anterior dorsal-fin rays intensely pigmented by dark chromatophores. Anal fin light gray, with dark chromatophores mainly concentrated on interradial membranes. Caudal fin light gray, with scattered dark chromatophores on membranes and rays. Pectoral and pelvic fins light gray, with scattered dark chromatophores on rays. Head darker dorsally than ventrally. Opercle and infraorbitals with few dark chromatophores on their surfaces. Premaxilla, anterior portion of maxilla, dentary, and lips with greatly concentrated dark chromatophores. Variations in color pattern between males and females described under sexual dimorphism.

Color in life. Based on specimens from CZUT-IC 7890. Dorsum of body iron greenish. Silvery or whitish midlateral stripe on body flanks extending from posterior region of humeral blotch to caudal peduncle; region ventral to this stripe with whitish tonality. Humeral blotch faint and vertically elongate. Dark blotch on caudal peduncle present, but relatively inconspicuous. Dorsal fin hyaline, presenting some portions showing greenish or yellowish iridescence; bases of five anterior dorsal-fin rays intensely pigmented by dark chromatophores. Anal fin grayish or hyaline, with tips of rays blackish. Caudal-fin hyaline, with distal margin yellowish and bases of lobes with somewhat greenish hue. Adipose fin orange. Pectoral fin grayish. Pelvic fin hyaline, somewhat grayish on its anterior portion. Head orange dorsally; golden and greenish on frontoparietal region and between nasal and mesethmoid. Premaxilla, anterior portion of maxilla, dentary, and lips with intense dark pigmentation, more concentrated on lower jaw than on upper jaw. Variations in color pattern between males and females described under sexual dimorphism.

Sexual dimorphism. Males with bony hooks on anal-, caudal-, pectoral-, and pelvic-fin rays. Caudal fin with short, slender, anterodorsally oriented hooks, especially on unbranched and branched portions of rays 14–18 ( Fig. 23 C). Pectoral fin with few minute bony hooks scattered on distal portion of outermost branched ray. Pelvic-fin rays with short, slender, anteroventrally oriented hooks on nearly entire length of each ray (except unbranched ray with one hook); hooks usually paired or one per segment. Anal fin with slender, anterolaterally placed hooks with broad bases; from 1 to 17 pairs per ray located on posteriormost unbranched ray and up to ten anterior branched rays (larger hooks on middle rays of this range). In adult males, posteriormost unbranched anal-fin ray and up to first 18 branched rays slightly longer than remaining rays, resulting in convex-shaped margin. In females, anal-fin rays gradually decreasing in length from anteriormost branched ray to posteriormost branched ray; anal-fin margin somewhat concave or straight. Anal-fin base of adult males slightly concave along its midlength, this base straight in females. Adult females with large, conical, externally developed urogenital papillae.

In life, males with dark pigmentation, sometimes whitish, orange or reddish, on distal portions of first two branched pectoral-fin rays. Alcohol specimens presenting brown, black, or whitish pigmentation ( Fig. 22 ). Females present light orange or yellowish hue on dorsal and anal fins. In life- or alcohol-preserved specimens, caudal peduncle blotch more intensely pigmented on middle caudal-fin rays of females than males. In males, posterior half of outermost branched pectoral-fin ray thicker, and highly branched distally. Distal portion of this ray splitting into two equal branches; each branch dividing into two smaller branches and each one of these splitting into nine or twelve branches, splitting further into minute branches to form fan-shaped structure (for picture of similar structure in G. t o r re s i, see Vanegas-Rios et al. 2013: fig 3). Fan-shaped structure formed by 28 (1), 30 (1), 32 (1), 38 (1), 40 (1), 42 (2), 46 (1), 48 (1), 50 (2), 53 (2), or 54 (1) minute branches. Females having this pectoral-fin ray splitting into two main branches and each one of these dividing into three or four pairs of branches, splitting further one more time. Often this ray with similar branching pattern of other branched rays.

Mature males with hypertrophied scale forming pouch on lower caudal-fin lobe and with ventral procurrent rays 2 and 3 forming spur-shaped structure. Scarce small aggregations of apparent glandular tissue located on caudal-fin rays and medially to pouch scale. First ventral procurrent ray with moderate concavity on its ventral margin ( Fig. 23 C). Second ventral procurrent ray somewhat longer than third ray, reaching posteriorly midlength of first ventral procurrent ray, and somewhat flattened in sagittal plane (especially on its posterior portion). Posterior portion of third ventral procurrent ray weakly developed laterally. Pouch scale with 16–38 radii, usually between caudal-fin rays 16 or 18 and third ventral procurrent ray. Posteroventral pouch-scale lobe of varied size, extending posteriorly along midlength or entire length (populations of San Jorge River basin) of third ventral procurrent ray ( Figs. 23 C–D). Dorsal surface of pouch scale attached via soft tissue (apparently connective) to caudal-fin rays 10 or 11 to 15–17. Posterior margin of pouch scale between caudal-fin rays 15 or 16 and third ventral procurrent ray. Four or five scales in vertical series situated ventral to terminal lateral-line scale, overlaying posterior portion of pouch scale. Dorsal margin of pouch scale having minute canal tube in two males. Dorsal portion of pouch scale sometimes with minute spiny patch. Females with large scale with 20 (1), 21 (5), 23 (1), 24 (1), or 25 (1) radii on lower caudal-fin lobe.

Mature males with gill gland, formed by fusion of 9 (1), 15 (1), 19 (4), 20 (2), 21 (5), 22 (7), 23 (3), 24 (5), 25 (1), or 26 (1) anterior gill filaments of ventral limb of first gill arch. Total number of ventral limb gill filaments 20 (1), 22 (1), 24 (7), 25 (4), 26 (6), 27 (5), 28 (4), 29 (1), or 30 (1). Gill-gland length 7.0–11.4 % SL (mean = 9.8 % SL). Regression comparisons of morphometric data by sex with more pronounced differences for pectoral fin to pelvic fin length and pelvic-fin length (both with higher values in males than in females) in larger specimens as function of SL ( Fig. 24 ).

Distribution. Gephyrocharax melanocheir is widespread along the Magdalena, San Jorge, Ranchería, and Caribbean basins of Colombia ( Fig. 1 ).

Remarks. The holotype of G. m e l a n o c h ei r (FMNH 56049) designated by Eigenmann (1912) was not examined. In the 1980s, this specimen was loaned to Stanley H. Weitzman (Weitzman & Fink 1985). Since 2011, the holotype and the rest of the loaned specimens have started to be returned to the corresponding institutions (Lisa Parmer and Richard P. Vari†, Pers. Commun.). Unfortunately, during this process, the holotype could not be found. In a visit to the fish collection at USNM in 2013, I also failed to locate the holotype which is effectively lost. Therefore, herein a neotype (FMNH 125572) is designated from the paratype series FMNH 69554 with the same locality of the lost holotype, following the article 75.3.4 and the recommendation 75A of the ICZN (1999). According to the collection records, this paratype lot was composed by nine specimens, but only eight specimens were found in the jar and, consequently, one specimen can be possibly lost. The neotype is a male in good conditions (except for the body coloration, which is faded), presenting most diagnostic characters of the species.

Gephyrocharax melanocheir was recorded by Taphorn & Lilyestrom (1984), Bonilla-Rivero & López-Rojas (2001), and Bonilla-Rivero et al. (2002) in the Lake Maracaibo basin and Meachiche River (Caribbean versant) in Venezuela. Later, Bonilla-Rivero & López-Rojas (2013) found some morphological discrepancies (not specified) between the specimens that they identified earlier as G. melanocheir in the Caribbean versant ( Bonilla-Rivero & López-Rojas 2001; Bonilla-Rivero et al. 2002) and the species description. They did not examine specimens from the type locality. The conclusion of these authors was that the Caribbean specimens might correspond to an undescribed species of the genus. Five poorly-preserved specimens (UF 23806) from the Caribbean versant were previously identified by Vanegas-Ríos et al. (2013) as G. melanocheir . These specimens were reexamined taking into consideration the aforementioned considerations of Bonilla-Rivero & López-Rojas (2013), and no evidence was found to either confirm them as G. melanocheir or support them as a new species. Due to the small sample analyzed, which lack of larger males, the specimens were excluded from the examined material. The distributional records of G. melanocheir from the Caribbean versant in Venezuela should be better investigated. The presence of the species in the Ranchería River basin is the only record confirmed of the species in the Caribbean versant.

Material examined. All specimens from Colombia: ANSP 84439*, 38.2 mm SL, Tolima, Honda, approximately 5°12'18.42"N 74°44'13.49"W 447 m a. s. l. CAR 73, 6, 31.0– 36.7 mm SL (2 ds, 32.8–35.4 mm SL), Atlántico, Ponedera, Magdalena River basin, Grande creek, approximately 10°39'0.95"N 74°45'47.88"W 16 m a. s. l. CAS 44292 (previously IU 12696a–f), 1 paratype, 33.4 mm SL (rad), [Tolima], Bernal creek, near Honda, approximately 5°12'14.89"N 74°45'44.32"W 455 m a. s. l. CAS 44293 (previously IU 12697a–c), 1 paratype, 35.4 mm SL (rad), Soplaviento, town on the Dique de Cartagena, between Cartagena and Calamar, approximately 10°19'60.00"N 75°6'0.00"W 2 m a. s. l. CZUT-IC 196, 3, 35.6–37.3 mm SL (2 c&s 35.6–36.0 mm SL), Tolima, Coello, Gualanday creek at police station of Gualanday, approximately 4°16'49.54"N 75°1'58.85"W 468 m a. s. l. CZUT-IC 615, 15, 19.6–36.3 mm SL, Tolima, Coello, Potrerilla creek, approximately 4°16'57.90"N 75°1'53.80"W 463 m a. s. l. ANSP 139153, 10, 23.6–30.5 mm SL, Magdalena, De Aguja creek, 40 km SE of Santa Marta, approximately 10°58'22.18"N 74°15'10.00"W 2 m a. s. l. CIUA 250, 4, 31.2–35.0 mm SL, Caldas-Antioquia, Magdalena River basin, La Miel River, approximately 5°46'47.39"N 74°41'2.97"W 154 m a. s. l. CIUA 1060, 15, 25.9–37.8 mm SL (2 c&s 34.2–36.7 mm SL), Caldas, Magdalena River basin, Casanguilas creek, tributary of Guarinó River, approximately 5°18'36.69"N 74°53'28.42"W 513 m a. s. l. CZUT-IC 933, 8, 31.3–34.8 mm SL, Tolima, Tetuán River system, Peralonso River, approximately 3°52'12.40"N 75°16'7.59"W 377 m a. s. l. CZUT-IC 1878, 1, 34.3 mm SL, Tolima, Alvarado River at the bridge, approximately 4°31'3.04"N 74°59'5.03"W 517 m a. s. l. CZUT-IC 2164, 3, 22.1–22.3 mm SL, La Guajira, Vereda Reposo, Aguja River, approximately 10°57'34.00"N 74°10'26.00"W 27 m a. s. l. CZUT-IC 2175, 4, 12.4–22.8 mm SL, César, César River, Vereda Puente Canoas, approximately 9°38'54.20"N 73°38'47.83"W 39 m a. s. l. CZUT-IC 2187, 1, 33.4 mm SL, Norte de Santander, San Alberto River, approximately 7°45'20.82"N 73°23'18.37"W 120 m a. s. l. CZUT-IC 2209, 2, 11.7–16.6 mm SL, Santander, Vereda Villa Eva, Lebrija River route to San Alberto, approximately 7°35'11.50"N 73°33'50.43"W 77 m a. s. l. CZUT-IC 2327, 2, 39.3–41.9 mm SL, Tolima, La Joya creek, Vereda Chorrillo, approximately 4°50'19.95"N 74°46'37.87"W 265 m a. s. l. CZUT-IC 2626, 10, Tolima, Ibagué, Vereda Chucuní, Totaré River, Manjarres creek, approximately 4°29'08"N 75°04'35"W 734 m a. s. l. CZUT-IC 2990, 2, 25.9–36.2 mm SL, Huila, Vereda Letían, Yaguará system, La Boa creek, approximately 2°39'32.58"N 75°30'57.27"W 570 m a. s. l. CZUT-IC 5757, 1, 33.0 mm SL, Cundinamarca, Negro River system, La Morena creek, approximately 5°22'0.01"N 74°33'25.02"W 366 m a. s. l. CZUT-IC 5858, 6, 19.1–34.4 mm SL, Huila, upper Magdalena River, Yaguilga creek, 2°13'25.00"N 75°44'11.98"W 876 m a. s. l. CZUT-IC 6139, 3, 35.4–39.2 mm SL, Huila, Garzón, upper Magdalena River, Guandinosa creek, 2°23'40.00"N 75°32'47.00"W 830 m a. s. l. CZUT-IC 7883, 14, 23.7–31.0 mm SL, Sucre, El Pital, San Jorge River basin, Caño Caracolí, approximately 8°34'4948"N 75°11'1399"W 21 m a. s. l. CZUT-IC 7890, 6, 28.8–31.4 mm SL (2 c&s 29.3–31.4 mm SL), Sucre, Buenavista, San Jorge River basin, Caño San Vicente a tributary of Montegrande River, approximately 8°39'24.60"N 75°15'0.59"W 42 m a. s. l. CZUT-IC 7896, 3, 28.3– 31.0 mm SL, Sucre, Buenavista, San Jorge River basin, Caño Montegrande a tributary of San Jorge River, approximately 8°39'8.86"N 75°12'19.86"W 30 m a. s. l. CZUT-IC 7997, 7, 27.7–33.8 mm SL, Sucre, San Marcos, small Caños about 200 m at the bridge of San Jorge River, approximately 8°38'48.63"N 75°5'59.1"W 19 m a. s. l. FMNH 69554, 7 paratypes (3 rad), 31.3–41.3 mm SL, same data of the neotype. FMNH 125572 (previously FMNH 69554), neotype of Gephyrocharax melanocheir by present designation, male, 32.6 mm SL (rad), [Tolima], Bernal creek near Honda, approximately 5°12'14.89"N 74°45'44.32"W 455 m a. s. l. IAvH-P 1, 35, 18.8–34.7 mm SL, La Guajira, River Ranchería basin, Tabaco creek, 11°9'0"N 72°34'0"W approximately 81 m a. s. l. IAvH-P 10378, 2, 34.3–39.1 mm SL, Antioquia, Puerto Triunfo, Magdalena River basin, La Isla creek, 5°53'0"N 74°44'49"W approximately 252 m a. s. l. IAvH-P 10379, 2, 26.9–29.6 mm SL, Antioquia, Puerto Triunfo, Magdalena River basin, Dos creeks creek, 5°54'13.1"N 74°44'30.2"W approximately 249 m a. s. l. ICNMHN 3182, 31, 22.1–34.9 mm SL, Boyacá, Puerto Romero, approximately 5°50'2.05"N 74°20'23.04"W 244 m a. s. l. ICNMHN 5299, 8, 22.5–37.5 mm SL, Boyacá, Puerto Boyacá, Serranías Las Quinchas, Magdalena River basin, La Colorada creek, approximately 5°49'45.08"N 74°20'54.96"W 244 m a. s. l. ICNMHN 6459, 70, 15.6–39.4 mm SL, Boyacá, Puerto Boyacá, Serranías Las Quinchas, Magdalena River basin, La Colorada creek, approximately 5°49'45.08"N 74°20'54.96"W 244 m a. s. l. ICNMHN 6472, 16, 23.0– 37.8 mm SL, Boyacá, Puerto Boyacá, Serranía de las Quinchas, La Fiebre creek, approximately 5°55'59.88"N 74°21'59.86"W 204 m a. s. l. ICNMHN 9657, 188, 26.7–37.2 mm SL, La Guajira, Distracción, Chorreras, El Cerrado, Ranchería River, approximately 10°54'46.18"N 72°53'39.95"W 213 m a. s. l. ICNMHN 11045, 1, 37.5 mm SL, Caldas, La Dorada, Magdalena River basin, bridge Victoria on route to Mariquita, Bocarná creek at confluence with Guarinó River, approximately 5°17'35.63"N 74°52'37.81"W 434 m a. s. l. ICNMHN 11527, 1, 37.1 mm SL, Caldas, La Dorada, Magdalena River basin, bridge Victoria on route to Mariquita, Bocarná creek at confluence with Guarinó River, approximately 5°16'45.73"N 74°52'46.23"W 428 m a. s. l. ICNMHN 11762, 5, 22.0– 36.2 mm SL, Caldas, La Dorada, Victoria, Villa del Río Farm, Guarinó River, Canguillas creek, approximately 5°18'50.62"N 74°53'53.98"W 513 m a. s. l. ICNMHN 13551, 3, 29.9–36.2 mm SL, Caldas, La Dorada, Victoria, Bocatoma, La Dorada aquaeduct, Guarinó River, approximately 5°18'37.58"N 74°54'45.11"W 618 m a. s. l. ICNMHN 15897, 1, 33.7 mm SL, Caldas, Norcasia, spillwat at La Miel River dam, approximately 5°33'35.74"N 74°53'26.23"W 444 m a. s. l. ICNMHN 16095, 1, 33.0 mm SL, Caldas, Norcasia, spillway in La Miel River dam, approximately 5°33'35.74"N 74°53'26.23"W 444 m a. s. l. ICNMHN 16198, 2, 31.7–36.9 mm SL, Caldas, La Dorada, Vereda La Habana, La Palmera Farm, La Miel River, approximately 5°38'30.67"N 74°47'5.01"W 190 m a. s. l. ICNMHN 16296, 6, 32.5– 37.4 mm SL, Caldas, La Dorada, Vereda La Habana, La Palmera Farm, approximately 5°38'30.67"N 74°47'5.01"W 190 m a. s. l. ICNMHN 17777, 7, 31.8–33.5 mm SL, Santander, El Carmen de Chucuri, Vereda El Topón, La Colorada River basin, Topón River, 6°45'45"N 73°34'56"W 276 m a. s. l. ICNMHN 17790, 12, 33.8–39.4 mm SL, Santander, El Carmen de Chucuri, Vereda El Topón, La Colorada River basin, Topon River at the bridge, 6°43'16"N 73°32'49"W 348 m a. s. l. ICNMHN 17817, 2, 30.7–31.4 mm SL, Santander, El Carmen de Chucuri, Vereda El Topon, La Colorada River basin, Topón River, 6°45'45"N 73°34'56"W 276 m a. s. l. ICNMHN 17826, 24, 20.6–38.1 mm SL, Santander, El Carmen de Chucuri, Vereda El Topón, La Colorada River basin, Topón River, small tributary, 6°45'46"N 73°34'57"W 265 m a. s. l. IMCN 3425, 5, 23.3–29.1 mm SL, Tolima, Honda, Magdalena River basin, Bernal creek, approximately 5°12'19.73"N 74°45'46.09"W 284 m a. s. l. IMCN 3458, 4, 34.6–36.2 mm SL, Tolima, Coello, Magdalena River basin, police station of Gualanday, Gualanday creek, approximately 4°18'17.50"N 75°2'0.01"W 561 m a. s. l. USNM 79209, 2 paratypes, 31.9–35.2 mm SL (2 rad), [Tolima], Bernal creek near Honda, approximately 5°12'14.89"N 74°45'44.32"W 455 m a. s. l.

Figures

FIGURE 1. Geographic distribution of Trans-Andean species of Gephyrocharax occurring west from eastern versant of Andean Cordillera in northwestern South America to southern Central America.

FIGURE 6. Tukey box plots of (A) number of maxillary teeth, (B) number of branched anal-fin rays, (C) number of branched pelvic-rays, and (D) number of lateral-line scales comparing among Gephyrocharax species. Number of specimens between brackets.

FIGURE 22. Gephyrocharax melanocheir, (A) male, neotype by present designation, 32, 6 mm SL, FMNH 125572; (B) female, paratype, 35.4 mm SL, CAS 44293; (C) male, 33.8 mm SL, CIUA 1060. Additional illustrations in Vanegas-Ríos et al. (2013: fig. 7). White arrow indicates pigmentation of outer branched pectoral-fin ray.

FIGURE 23. Gephyrocharax melanocheir, (A – B) upper and lower jaws, female, 35.6 mm SL, CZUT-IC 196, right side; (C – D) detail of lower caudal skeleton and associated pouch scale, male, 36.0 mm SL, CZUT-IC 196, left side. Anguloarticular: aa, dentary: dt, maxilla: mx, premaxilla: pm, pouch scale: ps, posteroventral lobe: pvl, principal caudal-fin ray: pcr, retroarticular: ra, spur-shaped structure: sss, ventral procurrent ray: vpr. Lateral view. Scale = 1 mm.

FIGURE 24. Regression plot by sex of (A) pectoral fin to pelvic fin length and (B) pelvic-fin length as function of SL for Gephyrocharax melanocheir (in A: females, r = 0.96, r 2 = 0.92, ɑ = 0, β = 0.209, n = 48; males, r = 0.69, r 2 = 0.48, ɑ = 0, β = 0.180, n = 39, in B: females, r = 0.94, r 2 = 0.88, ɑ = 0, β = 0.153, n = 48; males, r = 0.77, r 2 = 0.60, ɑ = 0, β = 0.177, n = 39).