Torres etal2012 Astyanacinus yariguies

A New Species of Astyanacinus (Characiformes: Characidae) from the Río Magdalena System, Colombia Author(s): Mauricio Torres-Mejia, Emilton Hernández, and Victoria Senechal Source: Copeia, 2012(3):501-506. 2012. Published By: The American Society of Ichthyologists and Herpetologists URL: http://www.bioone.org/doi/full/10.1643/CI-10-160

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Copeia 2012, No. 3, 501–506

A New Species of Astyanacinus (Characiformes: Characidae) from the Rı´o Magdalena System, Colombia Mauricio Torres-Mejia1,2, Emilton Herna´ndez3, and Victoria Senechal4 Astyanacinus yariguies, a new species, is described from Rı´o Cascajales, Rı´o Magdalena system, in the municipality of El Carmen, Santander, Colombia. This species is assigned to Astyanacinus because it exhibits all the diagnostic characters of the genus, including 25–36 branched anal-fin rays (28–30 in the new species), and a coloration pattern with two humeral spots, a midlateral stripe formed by anteriorly oriented chevron bars, and a horizontally oriented oval-shaped caudal spot that continues over the median caudal rays. Astyanacinus yariguies differs from its congeners by having 13 predorsal scales forming a regular series, six teeth in the maxilla, nine scale rows between the dorsal-fin origin and lateral line, and a relatively short snout length. This is the first report of the genus Astyanacinus from the Rı´o Magdalena basin. Se describe Astyanacinus yariguies, especie nueva, del Rı´o Cascajales, sistema del Rı´o Magdalena, municipio de El Carmen, Santander, Colombia. La nueva especie es asignada a Astyanacinus por presentar todos los caracteres diagno ´ sticos para e´ste ge´nero, incluyendo 25–30 radios ramificados en la aleta anal (28–30 en la nueva especie) y un patro ´ n de coloracio ´n con dos manchas humerales, una banda lateral media formada por un patro ´ n de marcas verticales con forma de a´ngulo, y una mancha caudal que continu ´ a sobre los radios medios de la aleta caudal. Astyanacinus yariguies se diferencia de sus conge´neres por tener 13 escamas dorsales formando una serie ordenada, seis dientes en la maxila, nueve hileras de escamas entre el origen de la aleta dorsal y la lı´nea lateral, y un hocico relativamente corto. E´ste es el primer reporte del ge´nero Astyanacinus para la cuenca del Rı´o Magdalena.

T

HE Rı´o Magdalena is the largest Trans-Andean river, and it basin is the most scientifically explored, populated, deforested, and polluted in Colombia (Galvis and Mojica, 2007). Despite extensive explorations of this drainage basin, its fish fauna has not been fully inventoried. Since the first ichthyological exploration of the Rı´o Magdalena in the 19th century, a steady flow of new species have been described (Steindachner, 1878; Eigenmann, 1912, 1913; Miles, 1947; Dahl, 1971). Currently, more than 210 fish species are native to this river (Maldonado-Ocampo et al., 2008). In December of 2009, while surveying a previously unexplored tributary of the Rı´o Magdalena, we captured a characin that exhibited coloration patterns not previously described for any fish of the basin. The species presented a midlateral stripe with anteriorly directed chevron-shaped marks. This pattern is not a novelty in Characidae. Other characins that exhibit that pattern are Astyanax leopoldi, ‘‘Astyanax orthodus,’’ ‘‘Astyanax superbus,’’ Astyanax villwocki, Astyanacinus moorii, Astyanacinus multidens, Creagrutus mucipu, Hyphessobrycon heliacus, H. bifasciatus, Jupiaba pinnata, Moenkhausia pittieri, Nematocharax venustus, and Tetragonopterus carvalhoi. Multiple characters indicate that the new species described herein differs from these species. However, assigning the new species to the generic rank was a challenge. Although diagnostic characters at the genus level distinguish the new species from most of the species listed above (Eigenmann, 1917; Weitzman et al., 1986; Zanata, 1997; Vari and Harold, 2001; Garcı´a-Alzate et al., 2010; Melo et al., 2011), the new species fits the current definition of two distinct genera, Astyanax and Astyanacinus. Astyanax is defined by a combination of characters that includes two rows of premaxillary teeth, five teeth on the 1

inner premaxillary tooth row, a complete lateral line, a caudal fin not partially covered with minute scales, and dentary composed of four large anterior teeth followed by abruptly smaller teeth (Eigenmann, 1921, 1927). However, the morphological characters disgnostic of Astyanax are highly variable among Characidae (Marinho and Lima, 2009), and the monophyly of Astyanax is dubious (Ornelas-Garcia et al., 2008; Mirande, 2009, 2010; Javonillo et al., 2010; Oliveira et al., 2011). Not surprisingly, species that fit the diagnosis of Astyanax have been assigned to other genera (e.g., Zanata, 1997). That is the case of the species described here, which fits in Astyanacinus, a genus split from Astyanax. Astyanacinus was originally defined by the following combination of characters (Eigenmann, 1907): two series of premaxillary teeth, absence of conical teeth in anterior area of dentary, setiform gill rakers, upper jaw longer than half of head length, posterior border of maxillary extending posteriorly beyond the preorbital, notched preorbital, around six maxillary teeth, pores on lateral line scales, and a relatively straight junction between maxilla and premaxilla. D’Agosta (2011) recently revised Astyanacinus, conducting a phylogenetic analysis on morphological characters, and increased the list of diagnostic characters of the genus. D’Agosta defines Astyanacinus by the following characters: a lower jaw larger than eye-orbit; three or more teeth in maxilla; the posterior margin of maxilla posterior to vertical from lateral ethmoid; the pterygiophore of first anal-fin ray pointing dorsally; the distance between postcleithrum one and two shorter than height of postcleithrum one; the presence of anteriorly directed chevron-shaped marks; the premaxilla with two rows of teeth, the inner one with five, and the outer one with four teeth; the dentary with four large anterior teeth, followed by multiple smaller teeth; the

Present address: Department of Biology, University of California, Riverside, California 92521; E-mail: [email protected]. Send reprint requests to this address. 2 Grupo de Estudios en Biodiversidad, Universidad Industrial de Santander, A.A. 678, Bucaramanga, Colombia. 3 Municipio de Mogotes, Santander, Colombia; E-mail: [email protected]. 4 Department of Cell Biology and Neuroscience, University of California, Riverside, California 92521; E-mail: [email protected]. Submitted: 7 October 2010. Accepted: 1 March 2012. Associate Editor: R. E. Reis. DOI: 10.1643/CI-10-160 F 2012 by the American Society of Ichthyologists and Herpetologists

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humeral region with two diffuse vertical humeral bars and one well-defined oval-shaped horizontal mark over the anterior vertical mark; a caudal spot horizontally elongated, extending toward distal border of central caudal-fin rays; a total number of anal-fin rays 25–36; dorsal-fin rays ii+9; the lack of hooks in fins; and pectoral-fin rays i+11–13. The new species described here is assigned to Astyanacinus because it displays all the characters diagnostic of the genus. We compare the new species against the seven currently recognized Astyanacinus species listed by D’Agosta (2011): ‘‘Astyanax’’ orthodus, ‘‘Astyanax’’ superbus, Astyanacinus moorii, Astyanacinus multidens, and three undescribed species from upper Amazon river tributaries (A. n. sp. Bolivia, A. n. sp. Peru, and A. n. sp. Purus). Given the currently uncertain relationship between species of Astyanacinus and the nonmonophyletic Astyanax, we also provide a diagnosis of the new species against species of Astyanax. MATERIALS AND METHODS Measurements are presented as percentages of the standard length (SL), except for subunits of the head, which are given as percentages of the head length. All measurements were recorded with digital calipers to the nearest 0.1 mm while viewing with the aid of a dissecting microscope. Morphometric and meristic data were taken following Fink and Weitzman (1974), with the addition of the following measurements: dorsal to adipose fin distance, from posterior terminus of dorsal-fin base to adipose fin anterior terminus; dorsal to caudal fin distance, from posterior terminus of dorsal-fin base to posterior border of hypural fan center; snout to anus distance, from tip of snout to anterior border of anus; head height measured at vertical running through the base of supraoccipital process, perpendicular to longitudinal body axis; postorbital head length measured as the minimum distance from posterior border of eye to posteriormost termination of bony opercle; gape width measured between junctions of maxilla and dentary bones at both sides of the mouth, with the mouth agape in all but seven paratypes; maxilla length measured along longitudinal axis. Vertebrae were counted and pelvic bone was measured from radiographs. Vertebral counts considered four elements for the Weberian apparatus and one centrum for the terminal element of the ural complex. In the description of meristic characters, the count values of the holotype are indicated with an asterisk and the number of specimens with a particular count are given in parentheses. The influence of body size on morphometric variables was evaluated by regression analysis of log-transformed data. The scaling factor (b) is obtained from exponential function of standard length (y 5 aSLb). After log-transformation, the relationship becomes linear (log y 5 b log SL+log a). The scaling factor was compared against the isometric prediction (b 5 1 for linear measurements; McHenry and Lauder, 2006) using t-tests. Institutional abbreviations follow Sabaj Pe´rez (2010). Astyanacinus yariguies, new species Figure 1, Table 1 Holotype.—UIST 1752, 61.9 mm SL, Colombia, Santander, El Carmen, Rı´o Magdalena system, Rı´o Cascajales system, Rı´o Sucio, under bridge on road from El Carmen to Vereda

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Fig. 1. Astyanacinus yariguies, new species, holotype, UIST 1752, 61.9 mm SL, Colombia, Santander, El Carmen, Rı´o Magdalena system, Rı´o Cascajales basin, Rı´o Sucio, under bridge on the road from El Carmen to Vereda Islandia.

Islandia, 6u41914.60N, 73u35915.10W, M. Torres-Mejia and E. Herna´ndez, 16 December 2009. Paratypes.—24, 36.1–77.4 mm SL. All collected by M. TorresMejia and E. Herna´ndez in Colombia, Santander, El Carmen, Rı´o Magdalena system, Rı´o Cascajales system. UIST 1753, 8, 48.3–77.4 mm SL, collected with the holotype, 16 December 2009; ICNMNH 17642, 10, 42.0–74.7 mm SL, Quebrada La Concordia at Vereda Islandia, 6u35922.30N, 73u34958.10W, 17 December 2009; UIST 1755, 6, 36.1–59.0 mm SL, Quebrada Agua Blanca, Can ˜ o Volador at Vereda Islandia, 6u38911.40N, 73u35922.20W, 17 December 2009. Non-type material.—2 CS, 59.1–63.0 mm SL. UIST 1753, collected with holotype. Diagnosis.—Astyanacinus yariguies is diagnosed from other Astyanacinus by having a regular predorsal scale series (vs. irregular in other congeners) with 13 scales (vs. 14 scales; 16 in Astyanacinus multidens and 15 in ‘‘A.’’ superbus), six teeth in maxilla (vs. five or less in ‘‘A.’’ orthodus, Astyanacinus moori, A. n. sp. Bolivia, A. n. sp. Peru, and A. n. sp Purus), nine scale rows between dorsal-fin origin and lateral line (vs. 7–8 in ‘‘A.’’ orthodus, ‘‘A.’’ superbus, A. n. sp. Purus, and A. n. sp. Peru), and a relatively short snout length (14.9–25.3 vs. 22-7–31.4 in A. moori, 24.4–30.1 in A. multidens, 24.4–30.99 in ‘‘A.’’ orthodus, 24.3–28.9 in ‘‘A.’’ superbus, 23.7–29.9 in A. n. sp. Bolivia, 26.4–30.9 in A. n. sp. Peru, and 26.1–28.4% of head length in A. n. sp. Purus). Astyanacinus yariguies is readily distinguished from other characins (except for ‘‘A.’’ orthodus) by its coloration pattern: two humeral spots, midlateral stripe formed by anteriorly oriented chevron bars, and horizontally oriented ovalshaped caudal spot that continues over median caudal rays and never reaches a vertical through the last anal-fin ray base. Besides its coloration pattern, Astyanacinus yariguies is distinguished from species of Astyanax by the combination of 6 teeth in maxilla, 13 scales in a regular predorsal series, 36–39 lateral line scales, 9 scale rows above and 7 scale rows below the lateral line, ii–iii 28–30 anal-fin rays, and a maximum body depth 35.4–43.2% of SL. Description.—Morphometric data presented in Table 1. Dorsal head profile convex from upper lip to vertical through middle of eye, and straight or slightly concave from latter point to tip of supraoccipital. Ventral head profile slightly convex to nearly straight. Snout and lower jaw tips blunt, lower jaw slightly longer than upper jaw. Premaxilla with two rows of teeth, maxilla and dentary with one row each (Fig. 2A). Dentary with symphyseal diastema as wide as half

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Table 1. Morphometric Data and Scaling for Holotype and 24 Paratypes of Astyanacinus yariguies, New Species. Standard length is expressed in millimeters. Values 1 to 18 are percentages of standard length and 19 to 25 are percentages of head length. Scaling indicated with slopes of regressions between standard length and each trait. P-values (isometry, b = 1): ns.0.05$*.0.01$**.0.001$***.

Morphometrics

Standard length 1. Depth at dorsal-fin origin 2. Snout to dorsal-fin origin 3. Length of base of dorsal fin 4. Base of dorsal fin to adipose fin 5. Base of dorsal fin to caudal-fin base 6. Snout to insertion of pelvic fin 7. Snout to anus 8. Snout to origin of anal fin 9. Length of pelvic bone 10. Length of base of anal fin 11. Length of caudal peduncle 12. Length of longest dorsal-fin ray 13. Length of longest pectoral-fin ray 14. Length of longest pelvic-fin ray 15. Length of longest anal-fin ray 16. Least depth of caudal peduncle 17. Head height 18. Head length 19. Snout length 20. Gape width 21. Maxilla length 22. Upper jaw length 23. Orbital diameter 24. Postorbital head length 25. Interorbital width

Holotype

Range

61.9 43.2 55.0 15.7 25.2 37.4 48.6 58.8 66.1 8.4 35.1 10.7 27.3 22.5 17.7 17.3 12.5 29.6 25.6 22.9 40.5 34.7 51.3 37.1 52.3 31.8

36.1–77.4 35.4–43.2 48.6–55.0 13.6–17.1 21.0–26.4 36.5–40.9 44.4–48.8 53.3–60.0 60.2–66.1 7.0–8.7 31.7–35.6 10.4–12.8 22.0–30.3 17.0–24.1 15.0–23.8 14.5–22.0 10.4–12.6 25.6–31.2 24.9–28.3 14.9–25.3 31.4–40.6 26.9–34.7 42.3–57.9 27.1–37.1 38.1–54.9 25.3–35.6

width of symphyseal tooth (Fig. 2B). Body moderately elongate and compressed. Greatest body depth immediately anterior to dorsal-fin origin. Profile convex from tip of supraoccipital to anterior of dorsal-fin base, with minor hump immediately posterior to supraoccipital process in most larger specimens. Dorsal-fin base straight to slightly convex, intruding body outline at its posterior terminus. Dorsal profile slightly convex from posterior base of dorsal fin to adipose fin, and slightly concave from posterior end of adipose fin to procurrent caudal-fin rays. Ventral body profile convex at abdomen with slight concavity at base of pelvic fins, straight to rather convex at base of anal, and concave at caudal peduncle. Dorsal-fin origin along vertical approximately at middle of standard length. Profile of expanded dorsal fin triangular, anterior branched ray longest and posterior ray shortest. Origin of adipose fin slightly anterior to vertical through posterior terminus of anal-fin base. Profile of distal adipose-fin border semicircular. Insertion of pectoral fin slightly anterior to vertical through posteriormost tip of bony opercle. Profile of pectoral fin triangular, tip of pectoral fin nearly extending to insertion of pelvic fin. Profile of extended pelvic fin triangular, shorter than profile of pectoral fin. Insertion of pelvic fin located anterior to vertical through dorsal-fin origin. Tip of longest pelvic-fin ray sometimes short of reaching origin of anal fin. Border of anal fin slightly concave. Caudal fin forked. Lateral line from supracleithrum to base of caudal fin, with dorsal curvature concave and more pronounced at ventral border of posterior humeral mark.

Scaling Mean ± SD 57.0 38.9 52.0 15.3 24.2 38.7 46.6 57.5 63.0 8.0 34.1 11.5 25.5 21.7 17.6 16.9 11.6 28.6 26.5 20.4 36.3 30.8 49.2 31.8 48.1 30.8

6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6

12.7 1.9 1.5 1.1 1.4 1.4 1.2 1.5 1.6 0.5 1.1 0.8 2.1 1.9 2.7 1.9 0.5 1.4 1.0 3.2 2.5 1.9 2.8 2.9 3.8 2.8

b (±SE) 1.0 1.0 0.8 1.2 1.0 1.0 1.0 1.0 0.1 1.0 0.9 0.7 1.2 0.8 0.6 1.1 1.0 0.9 1.4 1.0 1.0 0.9 0.6 1.2 1.3

6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6

0.04 0.03 0.05 0.04 0.03 0.02 0.02 0.02 0.05 0.03 0.06 0.04 0.07 0.04 0.05 0.04 0.04 0.02 0.09 0.05 0.06 0.05 0.07 0.04 0.04

ns ns ** *** ns ns ns ns *** ns * *** * ** *** ns ns *** *** ns ns ns *** *** ***

Teeth of premaxilla tricuspid except for second tooth of inner row with five cusps. Inner tooth row with five teeth, the first (symphyseal) tooth longest, second tooth horizontally widest, and fifth tooth smallest. Three medial teeth of inner row broader than teeth on outer tooth row. Outer row with four tricuspid teeth, first one longest and horizontally widest. Maxilla with six teeth, two pentacuspid, followed by two tricuspid, and last two unicuspid. Maxilla extends posteriorly to vertical through anterior margin of orbit, and slightly curved upward. Dentary with four large medial teeth and eight posterior small teeth that gradually decrease in size. First two medial teeth with seven cusps, third and fourth pentacuspid, fifth to tenth tricuspid, and last two unicuspid. Dorsal-fin rays ii 8 (1) or ii 9*(24). Anal-fin rays ii 29*(16), iii 28 (6), iii 29 (1), or iii 30 (2). Pelvic-fin rays i 7(1) or ii 7*(24). Pectoral-fin rays i 13*(19) or ii 11 (6). Caudal fin invariably with ten rays in upper lobe and nine in lower lobe. Fin-ray hooks absent. Scales in longitudinal series 36 (6), 37 (9), 38*(8), or 39 (2). Scales in transverse series from origin of dorsal fin to lateral line nine. Seven scales in transverse series from insertion of pelvic fin to lateral line. Thirteen predorsal scales, arranged in a regular series. Scales along middorsal line between posterior termination of base of dorsal fin and adipose fin 11 (4) or 12*(21). Vertebrae 35 (7), 36*(17), or 37 (1). Supraneurals invariably five. Coloration in alcohol.—Chromatophores denser at dorsal surface of body, making this region look overall dark brown.

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mark, extending to caudal-fin base and finishing as thick, horizontally oriented oval-shaped caudal spot that continues over median caudal rays. Caudal spot never reaching vertical through last anal-fin ray base. Area ventral of midlateral stripe, from vertical running through the anterior terminus of anal-fin base to caudal peduncle, with chromatophores that follow myosepta, forming pattern of posteriorly directed thin vertical chevron bars. Tissue overlaying anal-fin pterygiophores with chromatophores that follow underlying bones. Chromatophores line membranes between rays, densely dispersed in dorsal, adipose, and caudal fins, and more scattered in pectoral and pelvic fins. Scaling analysis.—Scaling analysis presented in Table 1. All morphometric variables increased with size (b . 0, P # 0.006 for all traits). Negative allometry (i.e., b , 1, larger individuals proportionally shorter) in lengths of head, orbital diameter, caudal peduncle, pelvic bone, base of dorsal fin, and longest dorsal-, pelvic-, and anal-fin rays. Positive allometry (i.e., b . 1, larger individuals proportionally longer) in distance from base of dorsal fin to adipose fin, length of longest pectoral-fin ray, snout length, postorbital head length, and interorbital width. Distribution.—Astyanacinus yariguies is known from tributaries of the Rı´o Cascajales, in Rı´o La Colorada basin, Rı´o Magdalena system, in the municipality of El Carmen, Santander, Colombia (Fig. 3). Rı´o Cascajales drains a region of the western slope of the Serranı´a de los Yariguı´es or Serranı´a de los Cobardes in the Cordillera Oriental of Colombia. Fig. 2. Astyanacinus yariguies, UIST 1753, non-type collected with holotype, 63.0 mm SL. (A) Left lateral view of jaws with teeth. (B) Frontal-view of dentary, showing symphyseal diastema. Scale bar 5 2.5 mm.

Density of chromatophores gradually decreases ventrally, revealing yellowish ground coloration of body. Ventral surface of head and abdomen unpigmented. Body pigmentation rather darker with size. Midlateral stripe located in third scale row above the lateral line, from posterior border of head to caudal spot. A faint layer of guanine overlaps midlateral stripe posterior to humeral marks. Midlateral stripe formed by pattern of 10–14 anteriorly directed chevron-shaped vertical marks, separated from each other by two to three scales. Chevron marks formed by deep pigmentation between myomeres. Chevron marks decrease in size and definition toward caudal peduncle. Chevron pattern of chromatophores more distinct on smaller specimens. Chevron marks of larger specimens with thin branching offshoots of pigment. Two parallel and vertically elongated humeral marks, anterior darker and longer than other one. Main body of both marks vary in shape, frequently resembling a number three in left-side view, rarely a crescent-shape or a diffuse form. Main body of both marks extends from slightly above midlateral stripe to lateral line. Ventral portion of anterior mark with narrow diagonal extension tapering toward opercle. Dorsal portion of both marks is diffuse dorsally. Anterior mark with darker area covering two scales below midlateral stripe. Region between marks with few scattered chromatophores. Very thin, black line of chromatophores originating behind second humeral

Remarks.—Astyanacinus is distributed from Atlantic drainages from Panama to Rı´o Paraguay, and Pacific drainages in Colombia. Astyanacinus yariguies has a distribution within the limits of other species of the genus, although it is the first species of this genus described from the Rı´o Magdalena. The congeners with the closest distribution to A. yariguies are ‘‘A.’’ orthodus and ‘‘A.’’ superbus. ‘‘Astyanax’’ orthodus has been reported from Pacific drainages in Colombia from the Rı´o Patı´a to the Rı´o San Juan, and from Atlantic drainages ranging from the Rı´o Atrato in Colombia to the Rı´o Sixaloa in Costa Rica (Bussing, 1998; D’Agosta, 2011; Ruiz-C. et al., 2011). ‘‘Astyanax’’ superbus has a spotty distribution in mountain streams of the Rı´o Orinoco. The disjointed distribution of these three species suggests that A. yariguies has been separated from ‘‘A.’’ orthodus for at least ,10 Mya, the time Rı´o Magdalena has been separated from the the Atrato-Pacific slope watersheds (Albert et al., 2006). Similarly, geological evidence suggests that A. yariguies and ‘‘A.’’ superbus have been isolated for at least ,12 Mya, the time of the initial rise of Eastern Cordillera (Albert et al., 2006). This time frame is greater than that required for allopatric speciation in other freshwater fishes (Near and Benard, 2004). Etymology.—The species name refers to the Yariguı´es, the indigenous group that inhabited an area that includes the watershed of Rı´o Cascajales (Rodrı´guez, 2001). They fiercely defended their pristine territory for more than 400 years, which likely contributed to the preservation of the species described here. The Yariguı´es finally succumbed to invasion and extermination in the mid-20th century (Ortiz, 2008). The species name is treated as a noun in apposition.

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Fig. 3. Map of southwestern Santander, Colombia, showing position of holotype locality (star), and the two other type localities (asterisks) of Astyanacinus yariguies, new species. Gray squares represent municipalities, and M at top left corner indicates main stem of Rı´o Magdalena.

MATERIAL EXAMINED Range of SL given in mm. Astyanax atratoensis: USNM 306566, 4 paratypes, photograph, x-ray, 66.1–103.4, Colombia, Truando´. Astyanax filiferus: all from Colombia, Santander, Rı´o Sogamoso basin, Sabana de Torres, Rı´o Sucio: UIST 1945, 1, 84.6; UIST 1946, 1, 79.9; UIST 1947, 1, 74.6; UIST 1948, 1, 67.7. Astyanax magdalenae: CIUA 1420, 4, 59.3–81.5, Colombia, Caldas, Norcası´a, Rı´o Manso; CIUA 1917, 1, 94.0, Colombia, Santander, Rı´o Sogamoso basin, Puerto Wilches, Puerto Cayumba; UIST 1430, 1, 89.37, Colombia, Santander, Piedecuesta, Rı´o Chicamocha at Puente Pescadero. Astyanax microlepis: USNM 79167, 4 paratypes, photograph, x-ray, 69.8–91.1, Colombia, Cartago. ‘‘Astyanax’’ orthodus: USNM 55655, 1 holotype, photograph, x-ray, 84.7, Colombia, Truando´; ICNMHN 33, 1, 57.2, Colombia, Rı´o Atrato basin, Rı´o Sucio; ICNMHN 128, 9, 46.6–94.4, Colombia, Choco´ , Pavarando´ , Rı´o Baudo´ ; ICNMHN 207, 3, 46.94–55.79, Colombia, Choco´, San Juan, Noanama´; ICNMHN 1599, 1, 107.8, Colombia, Choco´, Rı´o Atrato basin, Rı´o San Francisco de Icho, Quebrada Feliciana; ICNMHN 3071, 1, 62.8, Colombia, Antioquia, Uraba´, Rı´o Jurabo´ at intersection with Chigorodo´–Medellı´n road. Astyanax sp. 1: all from Colombia, Rı´o Manso, Caldas, Norcası´a: CIUA 1464, 5, 48.2.1–52.4; CIUA 1670, 4, 69.4– 94.5. ACKNOWLEDGMENTS Special thanks to the Herna´ndez family and Fundacio´n Luzmimi for their support. The Policı´a Nacional of El Carmen took extreme measures for our security. Procedures for obtaining permits and loan of specimens were facilitated by M. Ramı´rez-Pinilla and E. Mantilla (UIS). Thanks to O. Mejı´a (ICNMHN) and M. Jime´nez-Segura (CIUA) for facilitating access to fish collections. INCODER provided permission to collect the specimens that formed the basis of this study. Pictures of preserved specimens were taken with the collaboration of S. Lee and E. Geledzhyan. Radiographs were taken by C. Thacker and J. Seigel (LACM). Thanks to D. Reznick and members of his lab for their kind collaboration. MARC-U program (UCR) funded VS. We thank F. D’Agosta

for his comments and for facilitating access to his thesis. This paper greatly benefited from comments by R. Vari and A. Furness. LITERATURE CITED Albert, J. S., N. R. Lovejoy, and W. G. R. Crampton. 2006. Miocene tectonism and the separation of cis- and trans-Andean river basins: evidence from Neotropical fishes. Journal of South American Earth Sciences 21: 14–27. Bussing, W. A. 1998. Peces de las Aguas Continentales de Costa Rica. Editorial de la Universidad de Costa Rica, San Jose´, Costa Rica. D’Agosta, F. C. P. 2011. Taxonomia e relac¸o ˜ es filogene´ticas do geˆnero Astyanacinus Eigenmann, 1907 (Characiformes: Characidae). Departamento de Biologia, Programa de Po´s Graduac¸aˆo em Biologia Comparada. Vol. M.Sc. Universidade de Saˆo Paulo FFCLRP, Ribeiraˆo Preto, SP. Dahl, G. 1971. Los Peces del Norte de Colombia. Ministerio de Agricultura, Instituto de Desarrollo de los Recursos Naturales Renovables (Inderena), Talleres Litografia Arco, Bogota´. Eigenmann, C. H. 1907. Fowler’s ‘‘Heterognathous Fishes’’ with a note on the Stethaprioninae. The American Naturalist 41:767–772. Eigenmann, C. H. 1912. Some results of an ichthyological reconnaissance of Colombia, South America. Indiana University Studies 8:1–27. Eigenmann, C. H. 1913. Some results from an ichthyological reconnaissance of Colombia, South America. Part II. Indiana University Studies 18:1–31. Eigenmann, C. H. 1917. The American Characidae. Part I. Memoirs of the Museum of Comparative Zoology 43: 1–102. Eigenmann, C. H. 1921. The American Characidae. Part III. Memoirs of the Museum of Comparative Zoology 43: 208–310. Eigenmann, C. H. 1927. The American Characidae. Part IV. Memoirs of the Museum of Comparative Zoology 43: 311–428. Fink, W. L., and S. H. Weitzman. 1974. The so-called cheirodontin fishes of Central America with descriptions of two new species (Pisces: Characidae). Smithsonian Contributions to Zoology 172:1–46.

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Galvis, G., and J. I. Mojica. 2007. The Magdalena River fresh water fishes and fisheries. Aquatic Ecosystem Health & Management 10:127–139. Garcı´a-Alzate, C. A., C. Roma´n-Valencia, and D. C. Taphorn. 2010. A new species of Hyphessobrycon (Teleostei: Characiformes: Characidae) from the San Juan River drainage, Pacific versant of Colombia. Zootaxa 2349: 55–64. Javonillo, R., L. R. Malabarba, S. H. Weitzman, and J. R. Burns. 2010. Relationships among major lineages of characid fishes (Teleostei: Ostariophysi: Characiformes), based on molecular sequence data. Molecular Phylogenetics and Evolution 54:498–511. Maldonado-Ocampo, J., R. Vari, and S. Usma. 2008. Checklist of the freshwater fishes of Colombia. Biota Colombiana 9:143–237. Marinho, M. M. F., and F. C. T. Lima. 2009. Astyanax ajuricaba: a new species from the Amazon basin in Brazil (Characiformes: Characidae). Neotropical Ichthyology 7:169–174. McHenry, M. J., and G. V. Lauder. 2006. Ontogeny of form and function: locomotor morphology and drag in zebrafish (Danio rerio). Journal of Morphology 267:1099–1109. Melo, B. F., R. C. Benine, T. C. Mariguela, and C. Oliveira. 2011. A new species of Tetragonopterus Cuvier, 1816 (Characiformes: Characidae: Tetragonopterinae) from the rio Jari, Amapa´, northern Brazil. Neotropical Ichthyology 9:49–56. Miles, C. 1947. Los Peces del Rı´o Magdalena. Ministerio de la Economı´a Nacional, Seccio´n de Piscicultura, Pesca y Caza, Bogota´. Mirande, J. M. 2009. Weighted parsimony phylogeny of the family Characidae (Teleostei: Characiformes). Cladistics 25:574–613. Mirande, J. M. 2010. Phylogeny of the family Characidae (Teleostei: Characiformes): from characters to taxonomy. Neotropical Ichthyology 8:385–568. Near, T. J., and M. F. Benard. 2004. Rapid allopatric speciation in logperch darters (Percidae: Percina). Evolution 58:2798–2808. Oliveira, C., G. Avelino, K. Abe, T. Mariguela, R. Benine, G. Ortı´, R. Vari, and R. Corre ˆ a e Castro. 2011. Phylogenetic relationships within the speciose family

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Characidae (Teleostei: Ostariophysi: Characiformes) based on multilocus analysis and extensive ingroup sampling. BMC Evolutionary Biology 11:275. Ornelas-Garcia, C., O. Dominguez-Dominguez, and I. Doadrio. 2008. Evolutionary history of the fish genus Astyanax Baird & Girard (1854) (Actinopterygii, Characidae) in Mesoamerica reveals multiple morphological homoplasies. BMC Evolutionary Biology 8:340. Ortiz, A. P. 2008. Geo Von Lengerke, Constructor de Caminos. Divisio´n de Publicaciones, Universidad Industrial de Santander, Bucaramanga, Colombia. Rodrı´guez, J. 2001. Etnias y culturas en el medio ambiente de Colombia, p. 408–445. In: El Medio Ambiente en Colombia. P. Leyva (ed.). Instituto de Hidrologı´a, Meteorologı´a y Estudios Ambientales de Colombia, IDEAM, Bogota´. Ruiz-C, R. I., C. Roma´n-Valencia, B. E. Herrera-M, O. E. Pela´ez, and A. Ermakova-A. 2011. Variacio´n morfolo´gica de las especies de Astyanax, subge´nero Zygogaster (Teleostei, Characidae). Animal Biodiversity and Conservation 34:47–66. Sabaj Pe´rez, M. H. 2010. Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. American Society of Ichthyologists and Herpetologists, Washington, D.C. Steindachner, F. 1878. Zur Fischfauna des MagdaleneStromes. Denkschriften der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen Akademie deer Wissenschaften, Wien 39:19–78. Vari, R. P., and A. S. Harold. 2001. Phylogenetic study of the Neotropical fish genera Creagrutus Gu ¨ nther and Piabina Reinhardt (Teleostei: Ostariophysi: Characiformes), with a revision of the cis-Andean species. Smithsonian Contributions to Zoology 613:1–239. Weitzman, S. H., N. A. Menezes, and H. A. Britski. 1986. Nematocharax venustus, a new genus and species of fish from the Rio Jequitinhonha, Minas Gerais, Brazil. Proceedings of the Biological Society of Washington 99: 335–346. Zanata, A. M. 1997. Jupiaba, um novo geˆnero de Tetragonopterinae com osso pe´lvico em forma de espinho (Characidae, Characiformes). Iheringia, Se´rie Zoologia 83:99–136.