Taxonomic review of the Neotropical genus Gephyrocharax Eigenmann, 1912 (Characiformes, Characidae, Stevardiinae) Author Vanegas-Ríos, James A. text Zootaxa 2016 4100 1 journal volume 10.11646/zootaxa.4100.1.1 98731fd0-8af6-422f-b59e-98c05234dea5 1175-5326 261565 E69BBCC0-775F-4F5C-B125-B890DE0FF7FF Gephyrocharax valencia Eigenmann, 1920a ( Figs. 30–32 ) Gephyrocharax valencia Eigenmann, 1920a : 11 –12 [original description, holotype CAS 44297 [IU 15131], type locality: “Maracay, Lake Valencia off dock of Paper Mill” (= Venezuela : Aragua, Maracay, Lake Valencia , near dock of paper mill)]. Myers in Eigenmann & Myers, 1929 : 477 , 484 [key, redescribed, figures not cited]. Schultz, 1944 : 323 –324, table 20 [key, distributional data, compared with G. venezuelae ]. Weitzman & Fink, 1985 : 104 [examined material]. Bonilla & López, 1995 : 33 –45 [morphological and genetic study]. Burns, Weitzman, Grier & Menezes, 1995 : 133 –135 [spermatozoid morphology, evidence of insemination]. Machado-Allison, Chernoff, Royero-León, Mago-Leccia, Velázquez, Lasso, López-Rojas, Bonilla-Rivero, Provenzano & Silvera, 2000 : 16 [ Venezuela : Esequibo River basin, Cuyuni River]. Bonilla-Rivero & López-Rojas, 2001 : 1 –10 [morphologically and genetically compared with Venezuelan congeners]. Taphorn, 2003 : 61 , 141–142, figs. 42, 133–134 [key, morphological and ecological data]. Weitzman, 2003: 225 [catalogue]. Lasso, Lew, Taphorn, DoNascimiento, Lasso-Alcalá, Provenzano & Machado-Allison, 2004 : 152 [listed from Venezuela ]. Rodríguez-Olarte, Coronel, Taphorn & Amaro, 2007: 2 [ Venezuela : Caribbean versant, Tocuyo River]. Maldonado-Ocampo, Vari & Usma, 2008 : 181 [listed from Colombia ]. Arai, 2011 : 82 : [karyotype: 2n = 52]. Bonilla- Rivero & López-Rojas, 2013 : 487 –495, tables 1–4, figs. 1–3 [phylogeographically compared with Venezuelan congeners]. Vanegas-Ríos, Azpelicueta, Mirande & Gonzales, 2013 : 283 [examined material]. Phillip, Taphorn, Holm, Gilliam, Lamphere & López-Fernández, 2013 : 9 , 16, 28, table 1, fig. 38 [listed from Trinidad and Tobago , key]. Vanegas-Ríos & Phillip, 2013 : 1329 –1334, table 1, figs. 1–2 [ Trinidad and Tobago : south coast of island of Trinidad , Moriquite and Moruga Rivers; morphological and ecological data]. Thomaz, Arcila , Ortí & Malabarba, 2015 : 5 , table 2, Add. File 5 [comments on insemination strategy, phylogenetic relationships, placed in Stevardiinae , Stevardiini]. Gephyrocharax valenciae . Géry, 1977 : 352, unnumbered fig. on page 353 [color in life, name misspelled]. Thomaz, Arcila , Ortí & Malabarba, 2015 : 10, fig. 5, Add. File 2 [comments on insemination strategy, sampled specimens, name misspelled]. Diagnosis. Gephyrocharax valencia differs from most of its congeners by lacking the humeral blotch ( vs . presence of this blotch, except in G. caucanus , G. chocoensis , and G. m a r t a e ) and the terminal lateral line tube between caudal-fin rays 10 and 11 ( vs . presence of this tube, except in G. melanocheir , G. sinuensis , and G. t o r re s i ). Gephyrocharax valencia is also distinguished from G. atracaudatus , G. caucanus , G. chocoensis , G. intermedius , G. sinuensis , and G. venezuelae by the absence of a rhinosphenoid bone ( vs . presence of this bone). From G. caucanus , G. chocoensis , G. sinuensis , G. t o r re s i , and G. venezuelae the species is differentiated by the absence of an osseous lamella between the first and second basibranchials ( vs . presence of this lamella). Gephyrocharax valencia differs from G. chocoensis , G. m a r t a e , G. sinuensis , and G. t o r re s i by the fewer number of pouch-scale radii in adult males (13–21 vs . 22–39). The presence of a tendinous fiber posteriorly inserted from flexor dorsalis superior to caudalfin rays ( vs . two tendinous fibers) distinguishes G. v a l e n c i a from G. major , G. sinuensis , G. t o r res i , and G. venezuelae . In G. valencia , the anterior cartilaginous portion of symplectic bone does not reach the anterior margin of the quadrate-metapterigoid fenestra ( vs . anterior portion of this bone reaching or surpassing anterior margin of such fenestra), which also differentiates it from G. torresi and G. venezuelae . Description. Morphometric data in Table 8 . Largest male 34.2 mm SL, largest female 39.8 mm SL. Body laterally compressed, with maximum depth at vertical through pelvic-fin origin or midpoint between this origin and anal-fin origin. Dorsal profile of head straight from tip of supraoccipital spine. Dorsal profile of body 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. 30 ). Ventral profile of body convex from tip of dentary to vertical crossing pelvic-fin origin, straight from this point to anal-fin origin, slanting posterodorsally and straight or slightly curved from this point to caudal peduncle origin. Anterior fontanel present, rarely reduced to narrow opening by 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. TABLE 8. Morphometric data of holotype (A), paratypes, and other examined specimens of Gephyrocharax valencia . Ranges of males including holotype values. M = mean; SD = standard deviation.

Characters	Males	Females and unsexed juveniles
Holotype	n	Range	M	SD	n Range M SD
Standard length (mm)	27.5	113	22.1–34.2	28.0	2.5	221 16.4–39.8 27.4 4.3
Percentages of standard length:
Depth at dorsal-fin origin	32.3	104	25.6–33.1	29.4	1.7	181 23.8–33.2 29.2 2.0
Snout to dorsal-fin origin	62.1	104	61.3–69.6	64.8	1.6	180 60.4–69.7 65.2 1.6
Snout to pectoral-fin origin	27.6	104	25.4–31.0	27.7	1.1	180 24.1–31.5 27.5 1.2
Snout to pelvic-fin origin	44.2	104	41.5–48.6	45.0	1.4	180 43.0–49.6 45.8 1.3
Snout to anal-fin origin	55.7	104	54.7–61.3	58.1	1.6	179 54.9–61.7 59.0 1.3
Dorsal fin to pectoral fin length	44.9	104	43.9–50.9	47.1	1.4	179 40.9–51.2 47.2 1.6
Dorsal fin to adipose fin length	28.0	103	23.0–31.7	26.5	1.5	178 24.0–29.5 26.7 1.1
Dorsal fin to hypural complex length Eye to dorsal-fin origin	40.4 49.7	104 104	33.1–41.6 45.0–56.6	37.3 52.8	1.8 1.6	180 33.0–41.9 36.9 1.8 180 45.2–56.7 53.0 1.8
Pectoral fin to pelvic fin length	17.3	104	16.1–21.4	18.4	1.1	179 14.9–22.9 19.0 1.1
Pelvic fin to anal fin length	14.6	104	13.2–16.7	14.8	0.8	179 12.5–17.1 14.9 0.9
Dorsal-fin length	21.4	104	17.4–22.8	19.4	1.2	179 16.7–22.1 19.4 1.2
Dorsal-fin base length	10.7	104	8.2–11.6	10.1	0.7	179 8.3–11.7 9.9 0.6
Pectoral-fin length	24.5	103	22.3–28.4	24.5	1.0	178 20.1–28.5 24.4 1.3
Pelvic-fin length	16.9	104	13.3–17.5	15.5	1.0	179 11.2–18.7 14.2 0.9
Anal-fin length	20.5	104	16.3–21.9	19.0	1.2	179 15.2–22.5 19.0 1.3
Anal-fin base length	39.1	104	33.4–40.8	37.4	1.3	179 32.9–40.0 37.2 1.4
Caudal peduncle depth	11.6	104	8.9–13.3	10.7	0.7	179 7.7–11.4 9.7 0.7
Caudal peduncle length	11.3	104	7.2–15.3	10.1	1.9	179 6.2–13.4 9.2 1.8
Bony head length	25.0	104	22.0–27.4	24.4	1.0	181 22.1–27.3 24.3 1.1
Percentages of head length:
Snout length	27.9	104	20.2–31.9	27.0	2.7	181 20.5–31.4 26.7 2.4
Horizontal eye length	32.8	104	31.2–39.4	35.2	1.6	180 30.0–41.2 35.7 2.0
Postorbital head length	41.3	104	34.3–44.3	40.2	1.6	179 34.7–44.5 39.6 1.8
Least interorbital width	38.9	102	33.8–39.7	36.7	1.2	175 32.1–41.5 36.7 1.5
Upper jaw length	43.1	104	39.9–47.5	43.4	1.4	179 37.2–48.7 43.2 1.9

Mouth superior, lower jaw projecting somewhat anterior to tip of upper jaw. Premaxilla with two rows of teeth. Outer row with 2 (9), 3* (89), 4 (184), or 5 (3) teeth, usually tricuspid, sometimes conical or bicuspid. Inner row with 4 (30), 5* (248), or 6 (8) teeth; symphyseal tooth tetracuspid, rarely pentacuspid, remaining teeth tri- to pentacuspid. Usually posteriormost tooth with fewer number of cusp or conical. Maxilla rarely toothless (7), usually with 1* (268) or 2 (11) teeth, conical to tricuspid (rarely tetra- or pentacuspid) ( Fig. 31 A). Maxilla posteriorly reaching vertical through anterior one-third of eye. Dentary with 8 (5), 9 (18), 10 (41), 11 (86), 12 (48), 13* (21), 14 (10), 15 (2), or 16 (1) teeth; three anteriormost teeth larger, pentacuspid (sometimes anteriormost tooth tetracuspid), followed by one median-sized tooth tri- to pentacuspid, and 4 (5), 5 (18), 6 (41), 7 (86), 8 (48), 9* (21), 10 (10), 11 (2), or 12 (1) smaller conical to tricuspid teeth ( Fig. 31 B). Dorsal-fin rays ii,7 (6), 8* (285), or 9 (1). Nine* proximal pterygiophores on dorsal fin (6 rad, 10 c&s). Dorsalfin origin located at vertical between anal-fin rays 6–12. Adipose fin present (atypically absent in two specimens), its origin located at vertical through bases of two posteriormost anal-fin rays. Anal-fin rays iv (246) or v* (47), 23 (2), 24 (2), 25 (2), 26 (20), 27* (81), 28 (80), 29 (78), 30 (22), or 31 (5). Twenty-six to 31 proximal pterygiophores on anal fin (6 rad, 10 c&s). Anal-fin origin closer to origin of hypural joint than to snout tip. Pectoral-fin rays i,8 (20), 9* (210), or 10 (55). Pectoral-fin distal tip posteriorly reaching one-seventh to one-half of pelvic-fin length. Pelvic-fin rays i,6* (281) or 7 (3). Pelvic-fin origin located at vertical between pored lateral-line scales 7–10 and anterior to body midlength. Caudal fin forked with 10/9 principal rays in all specimens. Scales cycloid, with numerous radii along posterior margin. Lateral line complete, pored scales 36 (2), 37 (12), 38 (45), 39* (77), 40 (68), 41 (61), 42 (18), 43 (4), or 44 (1). Terminal lateral-line tube absent. Predorsal scales 17* (43), 18 (87), 19 (84), 20 (54), or 21 (4). Scale rows between dorsal fin and lateral line 5 (34), 6* (250), or 7 (3). Scale rows between lateral line and anal fin 4 (31), 5 (250), or 6* (6). Scale rows between lateral line and pelvic fin 4 (152) or 5* (129). Circumpeduncular scales 13 (21), 14 (164), 15 (48), or 16* (15). One row of 10 (2), 11 (6), 12 (23), 13 (32), 14* (52), 15 (38), 16 (18), 17 (20), 18 (9), 19 (2), or 20 (1) scales forming sheath along anal-fin base. Total number of vertebrae 38* (3), 39 (7), 40 (5), or 41 (1); 15* (12) or 16 (4) precaudal and 23* (3), 24 (8), or 25 (3) caudal (6 rad, 10 c&s). Gill-rakers on dorsal limb of first branchial arch 6 (171) or 7* (64); ventral limb with 10 (2), 11* (139), 12 (92), or 13 (1). Color in alcohol. Ground color pale yellowish or pale brown, darker along mid-dorsal line and slightly lighter ventrally. Minute dark chromatophores scattered over body, in minor proportion or absent on lateral and ventral regions of abdomen and ventral region of caudal peduncle. Dark midlateral stripe diffuse (silvery in specimens retaining guanine), extending from vertical through pelvic-fin origin or slightly anterior to this point to posterior region of caudal peduncle. Dark chromatophores present along myosepta between lateral line and upper portion of anal fin. Humeral blotch absent, not differentiated from midlateral stripe. Dark blotch on caudal peduncle, horizontally elongate, and extending from posterior region of peduncle across interradialis muscles to base of caudal-fin rays. Dorsal fin light gray or faintly yellowish, with scattered dark chromatophores on membranes and borders of rays. Anal fin light gray, with dark chromatophores more concentrated on interradial membranes than on rays. Caudal fin light gray or faintly yellowish, with concentrated dark chromatophores on membranes and borders of rays. Pectoral and pelvic fins light gray, with few dark chromatophores mainly on membranes. Head darker dorsally than ventrally. Scattered dark chromatophores on opercle and infraorbitals. Premaxilla, anterior portion of maxilla, dentary, and lips with scattered dark chromatophores. Variations in color pattern between males and females described under sexual dimorphism. Sexual dimorphism. Males with bony hooks on anal-, caudal-, and pelvic-fin rays. Caudal fin with short, slender, anterodorsally oriented hooks, especially on branched portions of rays 14–17 ( Fig. 31 C). Pelvic-fin rays (except unbranched ray) with short, slender, anteroventrally oriented hooks on nearly entire length of rays (sometimes absent or in minor proportion on innermost ray); hooks usually paired or one per segment. Anal fin with slender, anterolaterally placed hooks with broad bases; from 1 to 15 pairs per ray located on posteriormost unbranched ray and up to seven anterior branched rays (larger hooks on middle or posterior rays of this range). In adult males, usually posteriormost unbranched anal-fin ray and up to 15–17 anterior 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 or curved along its midlength, this base straight in females. Adult females with external, short, ventrally oriented urogenital papillae. Adult males having faint patch formed by scattered dark chromatophores extending ventrally from anterior region of urogenital pore to point immediately anterior to pelvic-fin origin. Females without concentrated chromatophores on this area or, if present, in minor proportion than on males. 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 concavity on its ventral margin ( Fig. 31 C). Second ventral procurrent ray somewhat longer than third ray, reaching posteriorly midlength of first ventral procurrent ray, and flattened in sagittal plane, especially on its posterior portion. Posterior portion of third ventral procurrent ray weakly developed laterally. Pouch scale with 13–21 radii , usually located between caudal-fin rays 17, 18, or 19 and third ventral procurrent ray. Posteroventral pouch-scale lobe of varied size, extending posteriorly along third ventral procurrent ray ( Figs. 31 C–D). Dorsal surface of pouch scale attached via soft tissue (apparently connective) to caudal-fin rays 10 or 11 to 14. Posterior margin of pouch scale between caudal-fin rays 15 or 16 and third ventral procurrent ray. Four scales in vertical series situated ventral to terminal lateral-line scale, overlaying posterior portion of pouch scale. Dorsal margin of pouch scale slightly straight or slightly concave. Females with large scale with 12 (1), 14 (2), 15 (4), 16 (2), 17 (2), 18 (3), 19 (1), or 20 (1) radii on lower caudal-fin lobe. FIGURE 30. Gephyrocharax valencia , (A) male, holotype, 27.5 mm SL, CAS 44297; (b) male paratype, 30.4 mm SL, CAS 44295; (C) female, 26.8 mm SL, MCZ 59390; (D) female, 26.0 mm SL, MCZ 59642; (E) female, INHS 31627, 30.5 mm SL. Photographed specimens representing some geographic groups analyzed using SPCA in text. Additional illustrations in Vanegas-Ríos & Phillip (2013: fig. 1) and Phillip et al . (2013: fig. 38). FIGURE 31. Gephyrocharax valencia , (A–B) upper and lower jaws, female, 32.7 mm SL, ANSP 134924; (C–D) detail of lower caudal skeleton and associated pouch scale, male, 27.5 mm SL, MBUCV 3675. 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, left side. Scale = 1 mm. FIGURE 32. Sheared principal components analysis of morphometric data of Gephyrocharax valencia comparing among species organized by geographic groups: (A) left plots showing first and second principal components (PC1 vs . PC2); (B) right plots showing second and third principal components (PC2 vs . PC3). Sheared principal component: H, principal component centered by groups: S. Percentage of explained variance beside each component. Mature males with gill gland ( holotype with damaged filaments), formed by fusion of 7 (1), 8 (2), 9 (4), 11 (8), 12 (4), 13 (5), 14 (8), 15 (14), 16 (4), 17 (4), 18 (4), or 22 (2) anterior gill filaments of ventral limb of first gill arch. Total number of ventral limb gill filaments 20 (3), 21 (4), 22 (7), 23 (14), 24 (13), 25 (6), 26 (2), 28 (1), or 30 (1). Gill-gland length 3.5–9.2 % SL (mean = 6.2 % SL). Regression comparisons of morphometric data by sex with major differences in pelvic-fin length (higher values in males than in females), but larger specimens not completely separating on SL axis. Distribution. Gephyrocharax valencia is distributed in the Orinoco River basin, coastal Caribbean drainages of Venezuela , and coastal drainages of the island of Trinidad ( Fig. 3 ). Ecological notes. Bonilla-Rivero & López-Rojas (1995) and Vanegas-Ríos & Phillip (2013) respectively provided data on the habitat of two Venezuelan populations (Lake Valencia and Tuy River) and two populations of the island of Trinidad (Moriquite and Moruga Rivers). Winemiller (1989) reported the presence of dermal protuberances on the lower lip associated to particular conditions of the habitat in the Apure River. Winemiller (1992) provided ecomorphological data. Remarks. Bonilla-Rivero & López-Rojas (1995) studied the population variation of G. v a l en c i a in the Lake Valencia and Tuy River basins. The same authors further analyzed the morphological differentiation of G. valencia regarding other Venezuelan species of the genus ( Bonilla-Rivero & López-Rojas 2001 ). These studies did not find morphological variations for G. v a l e n ci a populations, although the authors concluded that it can be morphometrically separated (but not completely) from the rest of the Venezuelan species. Bonilla-Rivero et al. (2002) reported some differences between G. v a l e n c i a and G. venezuelae based on a fewer number of examined specimens. Bonilla-Rivero & López-Rojas (2013) again analyzed G. valencia but this time with a genetic approach. They found that G. valencia has a genetic structure more defined than G. venezuelae , and additionally, proposed that the diversification of the genus in Venezuela may be explained by the changing in the course of the Orinoco River basin along the geological time ( Bonilla-Rivero & López-Rojas 2013 ). Finally, Bonilla-Rivero & López- Rojas (2013) concluded that G. v a l e n c i a is more related to a possibly undescribed species of Gephyrocharax than to G. venezuelae (this putative undescribed species has already been discussed under the remarks of G. melanocheir ). In the SPCA of the morphometric data, the examined specimens of G. v a l e n ci a were grouped by basins (Apure: Apure River basin; Caribe: some coastal drainages of the Caribbean versant; Cuyuni: Cuyuni River basin; Delta: drainages near the Delta of the Orinoco River basin; Meta: Meta River basin; Trinidad ; Morogu-Moriquite rivers basins in the island of Trinidad ; Uyare-Orocopiche: nearby drainages between Uyare and Orocopiche rivers basins; and Valencia : Lake Valencia basin). The SPCA did not indicate any significant difference characterizing the groups along the PC2 and PC3 ( Fig 32 ). However, one population of the Apure group (UF 80511) was slightly different from the rest of the groups. This population is almost completely distributed along the PC2 between the coordinates -0.3 and -0.1 and along the PC3 between the coordinates –0.05 and 0.10 ( Fig. 32 : note that the symbols of the Apure group are in these ranges) and also is marked by less concentrated cloud of individuals across the PC2 (variance = 7.42 %). This PC2 was more affected by the variation of the caudal peduncle length (-0.83) and the snout length (0.43), variables explaining almost all the separation among the individuals. Subsequent comparison of the population UF 80511 using the meristic and other measurements confirmed that this actually belongs to G. valencia , and its morphometric divergence from other populations across the PC2 should be considered only geographic variation of the species. Material examined. Colombia : ANSP 133981, 10, 29.4–34.4 mm SL (2 c&s 29.9–33.0 mm SL ), Meta, Meta River basin, La Venturosa creek, between La Balsa and Puerto López, 4°5'N 72°58'W approximately 186 m a. s. l. ANSP 139816, 8, 17.7–18.7 mm SL , Meta, Meta River basin, Mozambique Lake ( Mozambique farm), at its coastal line on north side, 3°58'N 73°4'W 206 m a. s. l. CZUT-IC 5346, 10, 23.5–30.2 mm SL , Casanare, Paz de Ariporo, Chire Viejo River, 6°01'20.0"N 71°23'4.5"W 150 m a. s. l. IAvH-P 4972, 10, 22.1–28.6 mm SL , Meta, Puerto López, Meta River basin, Negrito River, 4°7'0"N 72°57'0"W 172 m a. s. l. NRM 23841, 6, 20.0– 25.7 mm SL , Meta, Negro River, about 2 km S of Rincon de Pompeta town, approximately 4°1'36.01"N 73°21'48.99"W 256 m a. s. l. UF 23824, 7, 17.0– 32.3 mm SL , Meta, Meta-Guatiquía rivers basins, Caño Negros about 18.7 km on route from Villavicencio to Puerto Porfía, approximately 4°6'57.30"N 73°23'17.67"W 275 m a. s. l. UF 33410, 2, 30.6– 34.4 mm SL , Meta, Meta River basin, Guacavía River ENE of Villavicencio, approximately 4°17'24.24"N 73°30'51.06"W 449 m a. s. l. UF 80967, 5, 26.7–30.4 mm SL ; UF 80968, 11, 26.2–30.8 mm SL , Meta, Meta River basin, small creek about 70 km E of Villavicencio on route to Puerto López, approximately 4°4'24.81"N 72°59'16.55"W 185 m a. s. l. Venezuela : AMNH 77783, 4, 22.7–26.6 mm SL , Guárico, Orinoco River basin, Caño W of route from Calabozo to San Fernando and about 35 km S of Fundo Masaguaral (Caño Falcón), approximately 8°14’N 67°35’W 55 m a. s. l. AMNH 217653, 2, 24.6–25.8 mm SL , Delta Amacuro, Orinoco River, mainstream of coastal north of Tres Canos island, 129.7 nautic miles, approximately 8°39.6’N 61°59’W 8 m a. s. l. ANSP 133986, 25, 21.6–37.0 mm SL , Carabobo, Vigirima River (tributary of Guácara River), about 10 km NW of Guácara, where “dirt road” connect to unpaved route, approximately 10°24'N 67°55'W 253 m a. s. l. ANSP 134185, 20, 21.9–28.1 mm SL , Guárico, tributary of Guárico River, 3 km S of Calabozo on federal route 2, approximately 8°51’N 67°28’W 88 m a. s. l. ANSP 134924, 2 c&s, 24.5–32.7 mm SL , Carabobo, tributary of Lake Valencia basin, Caño Cambur 11 km S El Valencia , approximately 10°5'44.56"N 67°52'49.25"W 425 m a. s. l. ANSP 159869, 25, 21.9– 32.7 mm SL (2 ds 30.4–32.7 mm SL ), Bolívar, Caño at concrete bridge near Fundo Malama ranch (Sr. Biales), about 11 km N Sipao, approximately 7°33'N 65°23'W 53 m a. s. l. ANSP 159881, 50, 22.1–32.5 mm SL , Bolívar, river and flooded area about 15 km N of Maniapure on route between Caicara and Puerto Ayacucho, 6°55'27.87"N 66°33'32.44"W 72 m a. s. l. ANSP 161047 , 15, 19.3–28.7 mm SL , Bolívar, Agua Blanca River, from 100 to 600 m under bridge crossing Caicara-Bolívar route, approximately 7°50'N 63°51'18"W 71 m a. s. l. ANSP 165291, 7, 20.0– 26.3 mm SL , Apure, tributary of Arauca River, Caño El Yagua, approximately 7°30'0"N 68°20'0"W 74 m a. s. l. ANSP 168015, 10, 22.9–27.0 mm SL , Bolívar, Yuruari-Cuyuni basin, Miamo River, on Guasipati-El Miamo route about 20 km SW El Miamo, approximately 7°38'0"N 61°50'0"W 212 m a. s. l. AUM 5307, 104, 15.3–29.2 mm SL , Portuguesa, Portuguesa River basin, Caño Igues, Guanare-Guanarito route on km 60, 8°49'39.00"N 69°20'42.00"W approximately 110 m a. s. l. AUM 22776, 21, 21.2–30.5 mm SL , Portuguesa, Apure River basin, Portuguesa River, at the bridge on route 5, 9°4'54.48"N 69°40'53.72"W approximately 147 m a. s. l. CAS 44294 (previously IU 15129), 99 paratypes , 27.4–38.2 mm SL , [Carabobo], Lake Valencia basin, Isla del Burro, approximately 10°9'58.21"N 67°43'50.85"W 410 m a. s. l. CAS 44295, 1, 30.4 mm SL , Aragua, Maracay, Bue River. CAS 44297, holotype of Gephyrocharax valencia , 27.5 mm SL (rad), male, Aragua, Maracay, Lake Valencia , near dock of paper mill, approximately 10°13'0.85"N 67°37'11.82"W 448 m a. s. l. INHS 27775, 20, 24.4–30.6 mm SL , Barinas, Apure River basin, Caño Curito on route 5, 7°58.41'N , 71°0.5'W approximately 202 m a. s. l. INHS 28356, 20, 22.8–32.3 mm SL , Apure, Apure River basin, Orichuna River, 62 km E Elorza on route to La Victoria in El Charal, approximately 7°8'40.32"N 70°2'54.16"W 107 m a. s. l. INHS 28368, 20, 17.2–25.7 mm SL , Barinas, Apure River basin, Caño 7°16.26' N 71°5.20'W approximately 114 m a. s. l. INHS 29877, 16, 24.2– 32.6 mm SL , Barinas, Apure River basin, tributary of Suripa River, Yaure on route 5, 36 km NE from La Pedrera, Caño Yaure, 7°42.05'N 71°17.77'W approximately 174 m a. s. l. INHS 31385, 20, 20.2–28.0 mm SL , Aragua, Apure River basin, Guárico River, Barbacoas, 9°28'44"N 66°59'03"W approximately 170 m a. s. l. INHS 31627, 15, 20.7–30.5 mm SL , Bolívar, Cuyuni River basin, tributary of Yuruari, 3 km W of Guasipati on La Pastora route, 7°28'17"N 61°55'01"W approximately 176 m a. s. l. INHS 31720, 16, 23.0– 29.2 mm SL , Bolívar, INHS 31720, Bolívar, El Miamo, Yuruari River basin, Guanare River, approximately 7°38'13.32"N 61°46'41.97"W 207 m a. s. l. INHS 34668, 14, 18.8–28.2 mm SL , Barinas, Apure-Masparro rivers basins, Caipe River, 2 miles E of La Luz, approximately 8°24'31"N 69°48'19"W approximately 119 m a. s. l. INHS 60438, 8, 28.2–39.8 mm SL , Carabobo, Lake Valencia system, Caño La Cumara, 3 km N of San Diego, 10°16.55'N 67°56.21'W approximately 492 m a. s. l. INHS 69242, 20, 26.3–33.4 mm SL , Bolívar, Orinoco River basin, Pao River, Caño El Rodeo, route 19 crossing near El Báquiro, approximately 7°30'42.26"N 64°33'23.85"W 80 m a. s. l. INHS 69285, 20, 21.8–32.6 mm SL , Bolívar, Uyare-Orinoco basin, Suapire River, route 19 at the bridge 29 km E from Cuchivero River, approximately 7°29'51.95"N 65°39'38.22"W 72 m a. s. l. LACM 43138.017, 150, 20.4–30.1 mm SL , Delta Amaruco, Orinoco River, small drainage, Caño Araguao, 112 nautic miles upstream from El Mar de Buoy, 8°37'60.00"N 61°42'60.00"W 21 m a. s. l. LACM 43354.029, 80, 16.0– 26.8 mm SL , Bolívar, Orinoco River basin, Orocopiche River, about 15 km from mouth of Orinoco River, downstream from bridge on route 19, 8°2'60.00"N 63°40'0.00"W 27 m a. s. l. LACM 43355.008, 8, 16.4–21.5 mm SL , Orinoco River basin, Orocopiche River, about 15 km from mouth of Orinoco River, downstream from bridge on route 19, 8°2'60.00"N 63°40'0.00"W 27 m a. s. l. MBUCV 3675, 5, 26.2–39.1 mm SL (2 c&s 27.5–33.9 mm SL ), Miranda, Tuy River, isolated well of main course near Ocumare del Tuy, approximately 10°7'53.34"N 66°46'46.31"W 174 m a. s. l. MBUCV 8400, 5, 23.1–27.2 mm SL , Portuguesa, Ospino River, approximately 9°7'55.73"N 69°15'51.76"W 115 m a. s. l. MBUCV 15414, 5, 27.0– 31.3 mm SL , Anzoátegui, Neverí River, 100 m upstream at the bridge of San Diego, approximately 10°5'55.44"N 64°34'3.86"W 37 m a. s. l. MBUCV 21163, 5, 27.7–32.0 mm SL , Barinas, Anaro River, about 10 minutes of mouth of Suripá, approximately 7°49'4.51"N 70°17'57.38"W 110 m a. s. l. MBUCV 21928, 5, 24.5–26.9 mm SL , Guárico, Orituco River, Prestamo on route to Píritu, approximately 8°48'24.50"N 67°22'37.54"W 81 m a. s. l. MBUCV 35336, 5, 25.8–28.8 mm SL , Bolívar, Guaigura River, tributary of Aro River, approximately 7°36'15.87"N 64°7'36.07"W 86 m a. s. l. MCZ 59390*, 1 (rad), 26.8 mm SL , Portuguesa, Apure River basin, Caño Maraca at the bridge on route from Guanare to Guanarito, approximately 8°56'28.84"N 69°35'12.47"W 115 m a. s. l. MCZ 59642*, 1 (rad), 26.0 mm SL , Delta Amaruco, Orinoco River on coastal bay, 8°33'60.00"N 62°16'0.00"W 11 m a. s. l. MCZ 93507*, 1 (rad), 29.8 mm SL , Guárico, unknown locality, 8°56'16.56"N 67°25'38.90"W 86 m a. s. l. UF 80511, 45, 18.4–36.2 mm SL , Guárico, Caño E of Calabozo, isolated wells. USNM 257494, 2 c&s, 31.0– 32.0 mm SL , Guárico, Guárico River, 6 km SE of Calabozo, approximately 8°44'58.41"N 67°25'37.11"W 76 m a. s. l. Trinidad and Tobago : ROM 87531, 40, 21.3–26.9 mm SL , West islands, Trinidad island, Moriquite River, 10°7'0.08"N 61°17'34.01"W 16 m a. s. l. ROM 88770, 30, 19.0– 28.3 mm SL , West islands, Trinidad island, Victoria, Moruga River, Mayaro reserve, 10°7'2.42"N 61°12'14.58"W 21 m a. s. l. USNM 349208*, 2 (rad), 24.8– 28.3 mm SL , West islands, Trinidad island, Moruga, Moriquite River, town on Basse Terre, 10°07.904’N 61°15.436’W approximately 46 m a. s. l. UWIZM 2010.14.253, 5, 23.1–29.9 mm SL (2 c&s 25.2–28.0 mm SL ), West islands, Trinidad island, Moriquite River, approximately 10°6'51.54"N 61°17'39.78"W 24 m a. s. l.