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Astyanax pirapuan: a new characid species from the upper Rio Paraguay system, Mato Grosso, Central Brazil (Characiformes, Characidae) VICTOR A. TAGLIACOLLO1, RICARDO BRITZKE2, GABRIEL S. C. SILVA3 & RICARDO C. BENINE4 Laboratório de Biologia e Genética de Peixes, Departamento de Morfologia, IBB-UNESP Campus de Botucatu, Botucatu, SP, 18618-970, Brazil. E-mail:
[email protected];
[email protected];
[email protected];
[email protected]
Abstract A new characid fish is described herein from tributaries of the Rio Aricá-Mirim, Rio Cuiabá basin, Chapada dos Guimarães, Mato Grosso State, Brazil. Astyanax pirapuan better conforms to the A. scabripinnis species complex and it can be distinguished from other species of this complex by a combination of characters, including 8–9 gill rakers on the upper limb of first branchial arch, 0–1 tooth on the maxillary, 17 to 21 branched anal-fin rays, 35–37 perforated lateral line scales and two humeral spots. Key words: Neotropical, Astyanax scabripinnis, taxonomy
Resumo Um novo caracídeo é descrito de tributários do Rio Aricá-Mirim, bacia do Rio Cuiabá, Chapada dos Guimarães, Estado do Mato Grosso, Brasil. Astyanax pirapuan pertence ao complexo de espécies Astyanax scabripinnis e pode ser distinguido das demais espécies desse complexo pela combinação de caracteres, incluindo 8–9 rastros branquiais no ramo superior do primeiro arco branquial, 0–1 dente no maxilar, 17–21 raios ramificados na nadadeira anal, 35–37 escamas perfuradas na linha lateral e duas manchas umerais.
Introduction The genus Astyanax was originally proposed by Baird & Girard (1854) to include their new species Astyanax argentatus, a small tetra from Upper Nueces River, Texas, USA, currently considered by Lima et al. (2003) a junior synonym of A. mexicanus (De Fillipi, 1853). To date, the genus Astyanax comprises around 130 valid species (Eschmeyer, 2010), basically defined by the presence of two-teeth rows in the premaxilla with five teeth in the inner row, complete lateral line, and caudal fin not covered by a sheath of small scales (Eigenmann, 1917; 1921). According to various authors, the genus as it stands cannot be considered a monophyletic unit (e.g. Rosen, 1972; Weitzman & Malabarba, 1998). Recently, Mirande (2009) proposed a hypothesis of the phylogentic relationships of members of the family Characidae and pointed out that the few analyzed species of Astyanax form a monophyletic clade along with Bryconamericus scleroparius (Regan 1908), Hyphessobrycon anisitsi (Eigenmann 1907), H. bifasciatus Ellis 1911, H. luetkenii (Boulenger 1887), Markiana nigripinnis (Perugia 1891) and Psellogrammus kennedyi (Eigenmann 1903), which, due to its instability, was only named the Astyanax Clade. According to Mirande’s (2010) subsequent analysis, the Astyanax clade is supported only by the putatively synapomorphic presence of one or absence of maxillary teeth. This author further discussed that, according to his hypothesis, this clade would require a subfamilial category, but given the low taxon sampling and some variations observed under different searches, such an informal provisional name was employed, pending specific contributions. Thus, little light was effectively shed on the phylogenetic relationships of Astyanax. The Astyanax scabripinnis species complex was first proposed by Moreira-Filho & Bertollo (1991), who studied karyotypical and morphological features of the Astyanax scabripinnis subspecies sensu Eigenmann (1921).
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Accepted by M.R. de Carvalho: 15 Dec. 2010; published: 26 Jan. 2011
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Posteriorly, Bertaco & Lucena (2006) reviewed Eigenmann’s characters and proposed a more precise diagnosis for this complex. According to these authors, the Astyanax scabripinnis species complex is composed by: A. brachypterygium Bertaco & Malabarba, 2001; A. cremnobates Bertaco & Malabarba, 2001; A. intermedius Eigenmann, 1908; A. ita Almirón, Azpelicueta & Casciotta, 2002; A. jenynsii (Steindachner, 1877); A. laticeps (Cope, 1894); A. leonidas Azpelicueta & Casciotta, 2002; A. microschemos Bertaco & Lucena, 2006; A. obscurus (Hensel, 1870); A. ojiara Azpelicueta & Garcia, 2000; A. paranae Eigenmann, 1914; A. paris Azpelicueta, Almirón & Casciotta, 2002; A. rivularis (Lütken, 1875); A. scabripinnis (Jenyns, 1842); A. totae Haluch & Abilhoa, 2005 and A. troya Azpelicueta & Casciotta, 2002. It is not clear if those authors considered A. turmalinensis Triques, Vono & Caiafa, 2003 a member of such complex. In addition, five species of Astyanax described after the important paper of Bertaco & Lucena (2006) are currently assigned to A. scabripinnis species complex: A. burgerai Zanata & Camelier, 2009; A. courensis and A. goyanensis Bertaco, Carvalho & Jerep, 2010; A. jordanensis Alcaraz, Pavanelli & Bertaco, 2009; A. varzeae Abilhoa & Duboc, 2007. Representatives of the A. scabripinnis species complex were, until recently, only known from headwaters of coastal rivers of south and southeastern Brazil and from Rio Paraná and Rio Uruguay systems (see Bertaco & Lucena, 2006; 2010). Nowadays, exceptions for this distribution are A. burgerai, described from a coastal river from Bahia, Northeastern Brazil, and A. courensis and A. goyanensis described from the upper Rio Tocantins, Goiás, midwest Brazil. We describe herein a new characid which represents the first species of the Astyanax scabripinnis species complex from the Rio Paraguay basin.
Material and methods Counts and measurements followed Fink & Weitzman (1974) and Garutti & Britski (1997). Measurements were taken point to point with a caliper on the left side of specimens (precision of 0.1 mm). Values in the parentheses indicate the number of specimens with a particular count and an asterisk indicates values of the holotype. Counts of vertebrae, supraneurals, gill-rakers on the first branchial arch, and procurrent caudal-fin rays were taken from cleared and stained (c&s) specimens prepared according to Taylor & van Dyke (1985), previously operated for confirmation of sex by direct examination of the gonads under a stereomicroscope (after Crim & Glebe, 1990).Vertebral counts included the four vertebrae of the Weberian apparatus and the terminal centrum counted as single element. Comparative data were taken from original descriptions and/or directly examined material. Comparisons with Astyanax scabripinnis were based on Bertaco & Lucena (2006). Examined specimens are deposited in the Laboratório de Genética e Biologia de Peixes, Botucatu, SP (LBP); Museu de História Natural Capão da Imbuia (MHNCI), Museu Nacional, Rio de Janeiro, RJ (MNRJ), Museu de Zoologia da Universidade de São Paulo, São Paulo, SP (MZUSP), and Museu de Zoologia da Universidade Estadual de Londrina (MZUEL).
Astyanax pirapuan, new species (Fig. 1; Table 1) Holotype. MNRJ 37621, undetermined sex, 45.4 mm SL, Brazil: Chapada dos Guimarães, Mato Grosso State, Rio Aricá-Mirim, Rio Cuiabá basin, 15°46’04”S 55°30’44”W, 25 Nov 2005, N. G. Machado & W. R. C. Assunção. Paratypes. Brazil, Chapada dos Guimarães, Mato Grosso state: MNRJ 29315, 25, 27.6–65.2 mm SL, Rio Aricá-Mirim, Rio Cuiabá basin, 15°43’11”S 55°27’18”W, 25 Nov. 2005, N. G. Machado & W. R. C. Assunção. MNRJ 29357, 4, 35.6–51.0 mm SL, small stream of Rio Aricá-Mirim, Rio Cuiabá basin, 15°47’28”S 55°29’46”W, 27 Nov 2005, F. Modesto & A. Fiorentino. MNRJ 37622, 9, 24.4– 55.7 mm SL, same data as holotype. LBP 5662, 10, (2 c&s), 29.9–57.3 mm SL, Santo Antônio de Leverger, Rio Aricá-Mirim, 15°46’04”S 55°30’44”W, 07 august 2007, M. N. Mehanna & A. Ferreira. Non-type material (unadequated fixed material). LBP 5067, 2, 29.9–36.7 mm SL, LBP 5668, 5, 48.5–58.2 mm SL, Brazil, Mato Grosso State, Santo Antônio de Leverger, Rio Aricá-Mirim, 15°46’04”S 55°30’44”W, 08 august 2007, M. N. Mehanna & A. Ferreira. Diagnosis. Astyanax pirapuan belongs to the A. scabripinnis species complex and is distinguished from all species of this complex by the combination of the following characters: number of branched anal-fin rays (17–21 vs. 13–17 in A. jenynsii and A. brachypterygium; 13–16 in Astyanax goyanensis; 13–17 in A. jordanensis; 15–17 in NEW ASTYANAX FROM CENTRAL BRAZIL
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A. cremnobates, A. microschemos and A. courensis); teeth on maxilla (0–1 tricuspidate tooth vs. 1 heptacuspidate in A. ojiara; 2 in A. intermedius; 2–5, usually, 3 in A. leonidas and A. totae; 3–4 in A. paris); scale perforated (35– 37, usually 36 vs. 31–34 in A. burgerai; 38–40 in A. paranae; 37–39 in Astyanax obscurus); humeral spots (two vs. one in A. ita and A. rivularis); number of gill rakers on the upper limb of first branchial arch (8–9 vs. 6–7 in A. laticeps; 7 in A. varzeae). Yet, it is distinguished from A. troya through bony hooks (only on anal-fin vs. on pectoral, pelvic, anal and caudal fins in A. troya); from A. scabripinnis by number of circumpeduncular scales (12–14 vs. 16 in A. scabripinnis); from A. turmalinensis by series of scales between lateral line and pelvic-fin origin (4–5 vs. 5–6, usually 5, in A. turmalinensis), and number of teeth (0–1 tricuspidate tooth vs 1–2 tri-pentacuspidate teeth, usually 2, in A. turmalinensis).
FIGURE 1. Astyanax pirapuan, holotype, undetermined sex, MNRJ 37621, 45.4 mm SL, Brazil, Mato Grosso, Chapada dos Guimarães, Rio Aricá-Mirim, Rio Cuiabá basin.
Description. Morphometric data of holotype and paratypes are summarized in Table 1. Elongate and moderately compressed body. Greatest body depth usually immediately anterior to dorsal-fin origin. Dorsal profile of head between vertical through posterior nostril and tip of supraoccipital spine straight or slightly convex. Profile of body convex from tip of supraocciptal spine to base of last dorsal-fin ray, and slightly concave from that point to adipose-fin origin. Ventral profile of head convex. Ventral profile of body slightly convex from pectoral-fin insertion to anal-fin origin. Body profile along anal-fin base posterodorsally inclined. Caudal peduncle with dorsal and ventral profiles nearly straight. Snout rounded from margin of upper lip to vertical through anterior nostrils. Head relatively small. Mouth terminal. Upper jaw slightly longer than or equal to lower jaw. Maxilla short, not extending up to vertical through half of orbit, and aligned at angle of approximately 45 degrees relative to longitudinal body axis. Premaxilla with two tooth rows. Outer row with 3 (2), 4* (29) or 5 (9) tricuspid teeth with central cusp slightly longer. Inner row teeth 4 (6), 5* (32) or 6 (1) three-pentacusp teeth with central cusp twice longer and broader than other cusps. Maxilla with 1* (35) or 0 (3), tricuspid teeth. Dentary with 4* (44) anteriormost pentacuspidate larger teeth , followed by 5 (2) smaller tricuspid or conical teeth which are gradually decreasing in size; central cusp in all teeth are always larger than lateral ones and slightly reclined back (Fig. 2). Dorsal-fin rays ii (40), 8 (1), 9* (39). First unbranched ray smaller than second one. Dorsal-fin origin posterior to middle body; distal border smoothly rounded. Dorsal-fin pterygiophores 10 (2). Anal-fin rays iii, 18 (5), 19 (3), 20 (1), iv-17 (1) 18 (3), 19* (15), 20 (11), or 21 (1). Anal-fin origin situated posterior to vertical through base of last dorsal-fin ray. Pelvic-fin rays i, 7* (40). Pelvic-fin origin anterior to vertical line through dorsal-fin origin; distal border slightly convex tip not reaching anal-fin origin. Pectoral-fin i, 8 (1), 9 (7), 10 (11), 11 (12), or 12* (9). Caudal fin forked and unscaled, lobes similar in size, with 19 principal rays (2). Dorsal procurrent rays 8 (2) and ventral procurrent rays 10 (2). Lateral line complete. Perforated scales in longitudinal series 35 (16), 36* (20) or 37 (10). Scale rows between dorsal-fin origin and lateral line 6* (49); scale rows between lateral line and pelvic-fin origin 4 (38) or 5* (9). Scale rows around caudal peduncle 12 (7), 13* (35) or 14 (5). Predorsal scales 40* (11). Presence of the sheath scale of analfin base and axillary scale on pelvic fin origin. Precaudal vertebrae 17(2); caudal vertebrae 18 (1) or 19 (1). Supraneural 5 (2). Gill-rakers 11/8 (21), 12/8* (18), or 12/9 (1).
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TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. TABLE 1. Morphometrics data of holotype (MNRJ 37621) and paratypes (MNRJ 29315; MNRJ 29357; MNRJ 37622 LBP 5662) of Astyanax pirapuan. SD = Standard deviation. N = 49. Holotype
Paratypes Range
Mean
SD
45.4
22.0–65.2
41.3
-
Greatest depth
32.1
30.7–37.4
34.3
1.7
Snout to dorsal-fin origin
52.8
51.2–56.7
53.9
1.3
Snout to pectoral-fin origin
28.5
26.3–31.5
28.8
1.2
Snout to pelvic-fin origin
49.8
48.0–53.3
50.2
1.4
Snout to anal-fin origin
69.2
64.8–69.9
67.3
1.4
Caudal peduncle depth
11.8
10.9–12.5
11.9
0.4
Standard length (mm) Percents of standard length
Caudal peduncle length
11.1
9.3–13.2
11.4
0.9
Pectoral-fin length
21.5
17.9–21.7
20.2
0.8
Pelvic-fin length
16.4
14.4–17.6
16.1
0.8
Dorsal-fin length
22.6
21.7–27.1
24.2
1.5
Dorsal-fin base length
12.8
11.8–14.3
13.1
0.7
Anal-fin length
18.5
16.3–20.4
18.3
1.1
Anal-fin base length
23.3
21.1–25.3
23.6
0.9
Eye to dorsal-fin origin
40.4
37.8–42.8
39.9
1.1
Dorsal-fin origin to caudal-fin origin
50.6
48.4–53.4
50.4
1.2
Head length
27.8
27.0–30.5
28.4
0.8
Head depth
19.8
19.2–23.0
21.2
1.1
Snout length
23.7
18.8–24.9
22.0
1.6
Upper jaw length
28.0
24.5–30.3
27.4
1.7
Horizontal orbital diameter
31.4
30.9–37.3
34.2
1.8
Least interorbital width
34.5
31.7–39.2
35.6
1.5
Percents of head length
Color in alcohol. Overall ground color pale yellow, darker dorsally and yellowish ventrally. Dark chromatophores densely concentrated on dorsal surface of head from upper lip to supraoccipital spine. Dark chromatophores sparsely distributed over infraorbitals and two anterior thirds of maxilla. Opercle with sparse and large dark chromatophores. Ventral portion of head less pigmented, with scattered dark chromatophores more concentrated on anteromedian area. Dark chromatophores concentrated on distal margin of scales resulting in conspicuous reticulated pattern. Mid-dorsal region darker than flanks. Body with black, pigmented, midlateral stripe extending from second humeral mark to base of middle caudal fin, usually wider and more densely pigmented on caudal forming small caudal spot. Humeral region with two vertically-elongated spots. Anterior spot conspicuous, vertically elongated with upper portion broader and more evident than lower portion. Posterior humeral spot diffuse, with variable shape rounded, not ventrally surpassing lateral-line. Region between spots pale. Fins with dark chromatophores forming lines along rays extension; small chromatophores homogeneously distributed over membranes and borders of rays. Adipose fin with scattered small dark chromatophores. Sexual dimorphism. Males of Astyanax pirapuan are promptly recognized by the presence of a pair of bony hooks on the distal segments of the last unbranched and and the first eight branched anal fin ray. The bony hooks are aproximately the same size. No other secondary sexually dimorphic feature was identified. Etymology. The specific name pirapuan, from Tupi-Guarani, meaning “mountain fish” (pira = fish; apu = mountain) is an allusion to the geological characteristics of the type locality, Chapada dos Guimarães, Mato Grosso State, Brazil.
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FIGURE 2. Astyanax pirapuan, paratype, LBP 5662, 54.3 mm SL. Upper and lower jaws, left lateral view. Scale bar = 1 mm.
Distribution. Astyanax pirapuan is known from tributaries of Rio Aricá-Mirim, Rio Cuiabá basin, Chapada dos Guimarães, Mato Grosso State, Brazil (Fig. 3). Remarks. Astyanax pirapuan represents the first report of A. scabripinnis species complex in the Rio Paraguay basin. Recent description have demonstrated that species of this complex have restricted geographical distributions (e.g. Astyanax brachypterygium; A. cremnobates; A. ita; A. ojiara; A. paris; A. totae; A. troya; A. turmalinensis; A. goyanensis; A. courensis), which also seems to occur with A. pirapuan, as we have been collecting in many river basins and it has not been reported in other regions. Six species of Astyanax are known from the Rio Paraguay basin: A. abramis, A. asuncionensis, A. lineatus, A. marionae, A. paraguayensis and A. pellegrini. These species are distinguished from A. pirapuan since they do not present the diagnostic features of the Astyanax scabripinnis species complex sensu Bertaco & Lucena (2006). Besides, Astyanax pirapuan can be easily distinguished from A. asuncionensis, A. abramis and A. paraguayensis by the absence of a conspicuous oval-shaped spot in humeral area (vs. presence of an oval-shaped spot in the humeral area). Yet, it can be distinguished from A. lineatus, A. marionae and A. pellegrini by the number of anal fin-rays (21–24 vs. 26–28; 27– 29 and 41–45, respectively). Comparative material. Astyanax abramis: LBP 8422, 5, 39.17–53.11 mm, tributary of Rio Sepotuba, Tangará da Serra, Rio Paraguay basin, Mato Grosso State. Astyanax assuncionensis: LBP 8427, 10, 27.86–36.98 mm SL, Rio Sepotuba, Rio Paraguay basin, Tangará da Serra, Mato Grosso State. Astyanax biotae: MZUEL 4529, 1, 53.47 mm SL, stream Água Mole, Rio Paranapanema basin, Diamante do Norte, Paraná State. Astyanax brachypterygium: MZUSP 62713, paratypes, 10, 37.2–59.5 mm SL, arroio Água Branca, Rio Uruguay basin, Bom Jesus, Rio Grande do Sul State. Astyanax burgerai: MZUSP 101245, holotype, 1, 53.2 mm SL, córrego Água Boa on road between Floresta Azul and Almadina, tributary of small stream Pancadinha, itself a tributary of Rio Almada, Alma-
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dina, Bahia State. Astyanax cremnobates: MZUSP 62712, paratypes, 10, 37.5–64.9 mm SL, tributary of Rio Santa Cruz, Rio Jacu basin, São Francisco de Paula, Rio Grande do Sul State. Astyanax intermedius: MZUSP 47745, 1, 106.3 mm SL, tributary of Pedra Branca, Rio Paraíba do Sul basin, Cunha, São Paulo State. Astyanax jordanensis: MZUSP 99138, paratypes, 8, 42.7–76.1 mm SL, middle Rio Jacu, Rio Iguaçu basin, Candói, Paraná State. Astyanax lineatus: LBP 7636, 2, 38.09–43.39, mm SL, Rio Coxipó-Açu, Rio Paraguay basin, Chapada dos Guimarães, Mato Grosso State. Astyanax marionae: LBP 5830, 1, 50.37 mm, Rio Coxipó-Açu, Rio Paraguay basin, Cuiabá, Mato Grosso State. Astyanax paranae: CAS 22555, 1, holotype (photograph), Paraná State; MHNCI 8138, 10, 48.45–86.55 mm SL, Rio Lageado do Pitó, Rio Tibagi basin, Palmeira, Paraná State; MZUEL 1601, 3, 41.39–57.89 mm SL, tributary of Rio Lapé, Rio Tibagi basin, Fazenda das Rosas, Castro, Paraná State; MZUSP 55000, 6, 36.1– 54.1 mm SL, creek tributary of Rio Tibagi, upper Paraná basin, Fazenda das Rosas, Castro, Paraná State. Astyanax rivularis: MZUSP 39724, 20, 49.3–77.8 mm SL, córrego Guará, Rio São Francisco basin, Três Marias, Minas Gerais State. Astyanax aff. rivularis: MZUEL 5011, 3, 69.91–87.77 mm SL, Rio das Velhas, Rio São Francisco basin, São Bartolomeu, Minas Gerais State. Astyanax totae: MNRJ 28652, 10, 36.9–66.3 mm SL, Rio Cascata, tributary of Rio Tortuoso, Rio Iguaçu basin, Balsa Nova, Paraná State; MHNCI 10338, 15, 32.3–71.3 mm SL, Rio Cascata, tributary of Rio Tortuoso, Rio Iguaçu basin, Balsa Nova, Paraná State. Astyanax varzeae: MHNCI 11688, 1, 83.1 mm SL, Rio São João, Rio da Várzea headwaters, a tributary of Rio Negro, Rio Iguaçu basin, Tijucas do Sul, Paraná State. Astyanax xavante: MZUSP 100376, paratypes, 15, 18.8–59.6 mm SL, creek tributary to the córrego Avoadeira, inside the Parque Estadual da Serra Azul, Barra do Garças, Mato Grosso State.
FIGURE 3. Area showing sites of occurrence of Astyanax pirapuran (Brazil, Mato Grosso State, Chapada dos Guimarães, Rio Aricá-mirim, Rio Cuiabá basin, 1543’11”S 5527’18”W). Star: type locality, white circle: paratypes, square: Cuiabá, MT.
Acknowledgments We are grateful to Paulo A. Buckup and Marcelo Britto (MNRJ) for the loan of the material and curatorial assistance; Osvaldo T. Oyakawa (MZUSP) and Oscar A. Shibatta (MZUEL) for the loan of material. Vinicius Abilhoa provided morphometrics data from Astyanax paranae (MHNCI 8138). Carla S. Pavanelli (NUP) and Angela M. Zanata (UFBA) provided informations for A. turmalinensis. VAT and RB are financially supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, Proc. 132938/2009-0 and Proc. 132968/2009-6, respectively), and RCB is financially supported by the Programa Jovem Pesquisador UNESP and FUNDIBIO. NEW ASTYANAX FROM CENTRAL BRAZIL
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Literature cited Bertaco, V.A. & Lucena, C.A.S. (2006) Two new species of Astyanax (Ostariophysi: Characiformes: Characidae) from eastern Brazil, with a synopsis of the Astyanax scabripinnis species complex. Neotropical Ichthyology, 4(1), 53–60. Bertaco, V.A. & Lucena, C.A.S. (2010) Redescription of Astyanax obscurus (Hensel, 1870) and A. laticeps (Cope, 1894) (Teleostei: Characidae): two valid freshwater species originally described from rivers of Southern Brazil Neotropical Ichthyology, 8(1), 7–20. Crim, L.W. & Glebe, B.D. (1990) Reproduction, p. 529–547. In: Methods for fish biology. C. B. Schreck and P. B. Moyle (eds.). American Fisheries Society, Bethesda, MD. Eigenmann, C.H. (1917) The American Characidae. Memoirs of the Museum of Comparative Zoology, 43, 1–102. Eigenmann, C.H. (1921) The American Characidae. Memoirs of the Museum of Comparative Zoology, 43, 209–310. Eschmeyer, W. (2010) Catalog of Fishes. Electronic publication in “World Wide Web”. Available from: http://www.calacademy.org/research/ichthyology/catalog. (access on: Sep, 20, 2010). Fink, W.L. & Weitzman, S.H. (1974) The so-called Cheirodontin fishes of Central America with descriptions of two new species (Pisces: Characidae). Smithsonian Contributions to Zoology, 172, 1–46. Garutti, V. & Britski, H.A. (1997) Descrição de uma espécie nova de Astyanax (Teleostei, Characidae), com mancha umeral horizontalmente ovalada, da bacia do Rio Guaporé, Amazônia. Papéis Avulsos de Zoologia (São Paulo), 40(14), 217–229. Lima, F.C.T., Malabarba, L.R., Buckup, P.A., Silva, J.F.P., Vari, R.P., Harold, A., Benine, R., Oyakawa, O.T., Pavanelli, C.S., Menezes, N.A., Lucena, C.A.S., Malabarba, M.C. S.L., Lucena, Z.M.S., Reis, R.E., Langeani, F., Cassati, L., Bertaco, V.A., Moreira, C. & Lucinda, P.H.F. (2003) Genera Incertae Sedis in Characidae. In: Reis, R.E., Kullander, S.O. & Ferraris, C. (Ed.), Check List of Freshwater Fishes of South and Central America. Edipucrs, Porto Alegre, pp. 106–169. Mirande, J.M. (2009) Weighted parsimony phylogeny of the family Characidae (Teleostei: Characiformes). Cladistics, 25, 1– 40. Mirande, J.M. (2010) Phylogeny of the family Characidae (Teleostei: Characiformes): from characters to taxonomy. Neotropical Ichthyology, 8(3), 385–568. Moreira-Filho, O. & Bertollo, L.A.C. (1991) Astyanax scabripinnis (Pisces, Characidae): a species complex. Revista Brasileira de Genética, 14, 331–357. Rosen, D.E. (1972) Origin of the characid fish genus Bramocharax and a description of a second, more primitive, species in Guatemala. American Museum Novitates, 500, 1–21. Taylor, W.R. & van Dyke, G.C. (1985) Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium, 9, 107–119. Weitzman, S.H. & Malabarba, L.R. (1998) Perspectives about the phylogeny and classification of the Characidae (Teleostei: Characiformes). In: Malabarba, L.R., Reis, R.E., Vari, R.P., Lucena, Z.M.S. & Lucena, C.A.S (Eds), Phylogeny and Classification of Neotropical Fishes. Porto Alegre, Edipucrs, pp. 161–170.
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