Astyanax lacustris (L��tken, 1875) (Fig. 1; Table 1) Tetragonopterus lacustris L��tken, 1875: 131 (type locality: Lagoa Santa [rio das Velhas, drainage of rio S��o Francisco]. Tetragonopterus jacuhiensis Cope, 1894: 88 (type locality: Rio Grande do Sul, restricted to the rio Jacu��, Rio Grande do Sul, by Fowler, 1906: 435). [Syn. nov.] Astyanax (Poecilurichthys) bimaculatus paraguayensis Eigenmann, 1921: 256 (type locality: Paraguay [River] and upper Tocantins [River]). [Syn. nov.] Astyanax bimaculatus asuncionensis G��ry, 1972: 3 (name in substitution of Astyanax (Poecilurichthys) bimaculatusparaguayensis Eigenmann, 1921, preoccupied by Bertoniolus paraguayensis Fowler, 1918). [Syn. nov.] Astyanax altiparanae Garutti & Britski, 2000: 67 (type locality: rio Grande, Volta Grande Dam, Miguel��polis, S��o Paulo, Brazil). [Syn. nov.] Diagnosis. Astyanax lacustris is distinguished from other species of the genus Astyanax by presenting a horizontally oval, black humeral spot and two brown vertical bars situated in the humeral region, characteristics that include it in the Astyanax bimaculatus group. It is distinguished from A. argyromarginatus, A. clavitaeniatus, A. goyacensis, A. incaicus, A. novae, A. rupununi, A. saltor, A. siapae, A. unitaeniatus, and A. utiariti by the absence of a conspicuous midlateral black stripe extending above the lateral line, from the humeral spot or just behind it (from the second vertical bar), until the caudal peduncle, continuing along the median rays of the caudal fin; the midlateral stripe continually narrows forward (vs. midlateral black stripe present), and one black spot is present on the caudal peduncle (vs. spot absent). From the other species of the same group, A. lacustris differs from A. bimaculatus and A. validus by the absence of maxillary teeth (one tooth present on the left side of one of 237 examined specimens) (vs. presence). It further differs from A. validus by the absence of horizontal lines, forming a zigzag pattern on body (vs. presence); and from A. bimaculatus by presenting a markedly concave external surface of the second tooth of the internal series of the premaxilla, with misaligned cusps (vs. teeth having slightly concave surfaces with cusps almost aligned and on the same plane). Astyanax lacustris differs from A. abramis by a reduced number of perforated scales on lateral lines 30���39 (rarely 40 or 41) (vs. 42���48). See ���Note on other species of the Astyanax bimaculatus subgroup caudal peduncle spot��� below for A. maculisquamis, A. borealis, and A. vittatus, and ���Remarks��� on Astyanax lacustris for A.orbignyanus. Description. Morphometric data summarized in Table 1. Body compressed and high, with greatest body depth in around origin of pelvic fins. Dorsal profile of head straight or slightly concave from region above eye to supraoccipital spine. Dorsal body profile convex from tip of supraoccipital spine to dorsal-fin origin; straight from that point to adipose fin origin. Ventral profile of body convex from mandibular symphysis to anal fin origin and posterodorsally slanted along anal-fin base. Caudal peduncle deep, nearly straight on dorsal and ventral margins. Snout rounded from margin of upper lip to vertical through anterior nostrils. Head somewhat pointed anteriorly in lateral profile. Mouth terminal, jaw isognathous. Maxilla extending posteriorly to vertical through anterior margin of orbit. Maxilla widened anteroposteriorly. Two tooth rows in premaxilla; outer row with 4 (140) or 5 (3) tricuspid teeth, central cusp longer; inner row with five teeth, gradually decreasing in length from first to fifth; usually with five or six cusps on first tooth, five, six or seven cusps on second to fourth teeth and four or five cusps on fifth tooth; central cusp twice as long and broad as other cusps. Maxilla without teeth (one tricuspidate tooth present in the left side of a specimen���MCP 47854 - of 237 examined).Four anteriormost dentary large teeth with five, six or seven cusps; remaining four to seven (9 c&s)(4���12 in Garutti &Britski, 2000) teeth smaller, uni- to tricuspid (Fig. 2). Dorsal-fin rays iii (9 c&s) (first very small), 8 (6), 9 (168) or 10 (33); second unbranched ray approximately half length of third ray. Distal margin of dorsal fin nearly straight to slightly convex. Dorsal fin origin slightly behind middle of SL. Origin of adipose fin at vertical through base of sixth, seventh or eighth last anal-fin rays. Anal-fin rays iv(6 c&s) or v(5 c&s), 21 (5), 22 (7), 23 (23), 24 (39), 25 (39), 26 (62), 27 (40), 28 (36), 29 (8), 30 (4) or 31 (1) (up to 34 in Garutti& Britski, 2000). Anal-fin profile smoothly concave. Anal fin origin posterior to vertical through base of last dorsal-fin ray. Pectoral-fin rays i, 11 (108), 12 (42), 13 (1), or 15 (2). Pectoral-fin tip reaching or slightly surpassing pelvic-fin insertion; not reaching in specimens smaller than 11.5 mm SL. Pelvic-fin rays i, 7. Pelvic-fin origin anterior to vertical through dorsal-fin origin. Pelvic-fin tip does not reach or barely reaches anal-fin origin. Caudal fin forked, lobes similar in size, with i+ 17 +i principal rays, no scales in the lobes. Dorsal procurrent caudalfin rays 7 (1), 9 (1), 10 (2), 11 (2) or 12 (2), and ventral 7 (1), 8 (1), 9 (4) or 10 (2) (8 c&s). Lateral line complete, with 32 (1), 33 (26), 34 (38), 35 (60), 36 (65), 37 (41), 38 (10), 39 (9) or 40 (2) scales (up to 41 in Garutti & Britski, 2000). Scale rows between dorsal-fin origin and lateral line 6 (103), 7 (121), 8 (26) or 9 (2); scale rows between lateral line and pelvic-fin origin 4 (1), 5 (88), 6 (115), 7 (46), 8 (13) or 9 (2); scale rows between lateral line and anal-fin origin 5 (4), 6 (36), 7 (66), 8 (73), 9 (74) or 10 (7). Scale rows around caudal peduncle 11 (1), 12 (8), 13 (25), 14 (65), 15 (66), 16 (60), 17 (15), 18 (12) or 19 (1). Precaudal vertebrae 16 (4), 17 (2) or 18 (1); caudal vertebrae 18 (5), 19 (1) or 20 (1); total vertebrae 34 (2), 35 (3) or 36 (2). Supraneurals 4 (5)or 5 (2). Gillrakers at first arch: upper branch 6 (7), 7 (10), 8 (47), 9 (104), 10 (75) or 11 (5); lower branch 12 (17), 13 (94), 14 (65), 15 (50), 16 (19), 17 (4) or 18 (1); total number 20 (6), 21 (29), 22 (46), 23 (71), 24 (52), 25 (33) or 26 (10). Color in alcohol. Dorsal and dorsolateral portions of head and body dark brown. A dark oval humeral spot horizontally placed, followed by two vertical bars tenuously darkened; one exceeds the oval spot and the other remains two or three scales from it. Dark spot on caudal peduncle conspicuous, extending over median rays of caudal fin; anteriorly, it can reach the vertical through the origin of adipose fin.Body with dark spots at the posterior region of scales forming 2 to 4 horizontal stripes above lateral line and 2 to 5 below (Figs 3 A, B) (see Remarks). For color of living or freshly collected specimens see Desordi et al. (2011), Venere & Garutti (2011), Malabarba et al.(2013), Serra et al.(2014), and Gra��a & Pavanelli (2007). Sexual dimorphism. Male specimenswith hooks on anal and ventral fins. Anal fin with hooks from last unbranched ray (can be absent) until the first five, or until the tenth branched ray; one pair per segment. Hooks are present on all branched rays of the pelvic fin. Gill fusion glands were not found macroscopically on the first gill arch. T��ran et al. (2014) have found glandular cells but not fusion on the gill filaments in A. asuncionensis (= A. lacustris). Geographic variation. Variation of meristic characters showed ample overlapping after comparison between the drainages. Therefore, they were not useful in proposing diagnoses according to our criteria. This does not mean that we have not found statistically significant differences in the analysis undertaken. For example, Dunn��s method indicates significant differences in the number of total gill rakers and lower branch gill rakers between the populations of rio S��o Francisco and the other three populations from Paraguai, Paran�� and SLPUR (Figs. 4 A, B). The last count was significantly different between the rios Paran�� and Paraguai and SLPUR (Fig. 4 B). The number of gill rakers on the upper branch was significantly different between populations of the Paran�� and Paraguai vs. the S��o Francisco and SLPUR (Fig. 4 C). Significant differences were found in the number of branched anal fin rays between the S��o Francisco and Paraguai and SLPUR (Fig. 4 D). The number of perforated scales of the lateral line and of scales around the caudal peduncle was significantly different between the rio S��o Francisco populations and those of the Paran�� and Paraguai. The same occurred between the populations of SLPUR and the Paraguai (Figs. 4 D, E). The number of scales around the caudal peduncle was significantly different between SLPUR and Paran�� populations (Fig. 4 F). Significant differences were found in the number of scales from the lateral line and pelvic fin origin between S��o Francisco and Paraguai and SLPUR; the same occurred between the Paran�� and Paraguai and SLPUR (Fig. 5 A). The scale count from lateral line to anal fin origin was significantly different between therios S��o Francisco, Paraguai and Paran��, and SLPUR; the same count was different between the rios Paraguai and Paran�� (Fig. 5 B). Significant differences were found in the number of scales from dorsal fin to lateral line between the rio Paraguai and the other three populations (Fig. 5 C). Morphometric data analyzed did not indicate diagnostic differences (Tab. 1), and in an exploratory test PCA did not reveal discriminate populations at all (Fig. 5 D).In PCA, the three first axes explained 96.26 % variation of the data, but the scree plot between axes indicated that only the first two are informative. The axis PC 1 corresponds to the axis of size with all variables showing positive loadings. Axis PC 2, however, showed the population of the Paran�� partially discriminated from populations of the rios S��o Francisco and Paraguai, and SLPUR. The most influential variables in this discrimination in the second axis (explaining 2.892 % of variance) were diameter of the eye and interorbital width on the positive side, and pre-anal distance on the negative side. The drainages ��� the rios S��o Francisco, Paraguai and Sistema da Laguna dos Patos��� did not show discrimination between them. FIGURE 4. Astyanax lacustris populations by river drainages.(A) Number of total gill rakers on the first gill arch in. (B) Number of gill rakers on lower branch of first gill arch. (C) Number of gill rakers on upper branch of first gill arch. (D)Number of branched anal-fin rays. (E) Number of perfored scales of lateral line. (F) Number of scales around caudal peduncle. Tukey box plots: the middle line is the median and lateral lines of the box are 25 th and 75 th respectively. FIGURE 5. Astyanax lacustris populations by river drainages (A) Number of scales between pelvic-fin and lateral line in. (B) Number of scales rows between anal-fin and lateral line. (C) Number of scales rows between dorsal-fin and lateral line. Tukey box plots: the lateral lines of the box are 25 th and 75 th respectively. (D) Projection of individual scores in the space of the first and second Principal Component axis for the populations of Astyanax lacustris. In specimens, principally from the rio S��o Francisco drainage, anterior prolongation of caudal spot can be more tapered (contrary to truncated, a more common form in the other drainages), reaching the vertical through nearly half the distance between origin of adipose fin and end of dorsal-fin base (Figs. 1 A, B). This prolongation, however, sometimes is difficult to visualize, both in its form and scope, suggesting that it may be related to fixation (Fig. 6 A). Distribution. Rio La Plata and rioS��o Francisco drainages, Sistema da Laguna dos Patos and rio Tramanda�� drainage, and according to Garutti & Langeani (2009),in rio Araguaia (Tocantins-Araguaia drainage) (Fig. 7 A). Remarks. Tetragonopterus lacustris was described by L��tken based on specimens captured in Santa Lagoon, rio S��o Francisco, and taken by Reinhardt to the Zoological Museum at the University of Copenhagen. L��tken did not mention the number of specimens examined, but according to M. Krag (pers. comm.), collection manager of this museum, there are 25 specimens catalogued in that museum. Other syntypes are in the Museum f��r Naturkunde, Berlin, Mus��e National d'Histoire Naturelle, Paris, and in the Staatliches Museum f��r Naturkunde, Stuttgart. Photos of all specimens were examined and confirmed as belonging to the A. bimaculatus subgroup with caudal peduncle spot (see type material examined)(Figs. 1 C, D). Astyanax paraguayensis was described by Fowler (1918) as type species of the new genus Bertoniolus. G��ry (1972) synonymized Bertoniolus with Astyanax. To avoid a homonymy with Astyanax (Poecilurichthys) bimaculatus paraguayensis Eigenmann, 1921, G��ry proposed for this species the substitute name A. bimaculatus asuncionensis, maintaining as such the name A. paraguayensis for the Fowler species. Extensive series of specimens were analyzed from the rios La Plata and S��o Francisco drainages and SLP, as well as the syntypes (Fig. 6 B). In light of this analysis, and considering that the only characteristic mentioned by Eigenmann to distinguish Astyanax bimaculatus paraguayensis from the others subspecies recognized by him (lacustris, novae, vittatus, paraguayensis, and borealis) was ���rows of spots along the centers of the scales���, a pattern widely observed in the populations from the studied area (Figs. 6 C), a synonymization is proposed for the name A. bimaculatus asuncionensis with A. lacustris, the oldest name (see details on coloration further below). The diagnostic characters of A. altiparanae (Fig. 8 A) presented by Garutti & Britski (2000), are: ��� 33 to 41 scales on the lateral line, 10 to 17 scales on the transversal line, 22 to 34 rays on the anal fin, lower portion of body (when divided by the horizontal line that passes through the oral cleft) greater than the upper portion, 4 to 12 (frequently 6 to 8) small teeth on the dentary; large dentary teeth with five, six, or seven cusps; latero-ventral and ventral region of the trunk silvery, absence of maxillary tooth and black lateral band���. While comparing the new species with A. lacustris, the authors affirm that it possesses ���the same uniform coloration as altiparanae on the ventral flank, differing from it by other characters, among which are the approximately equal height between the upper and lower portions of the horizontal line that passes through the oral cleft (vs. lower portion higher than the upper in A. altiparanae), the large teeth with five or six cusps on the dentary bone (vs. five, six, or seven cusps in A. altiparanae), and the lesser number of small teeth on the dentary (3 to 6) (vs. 4���12, usually 6���8 teeth in A. altiparanae)���. The last two characters overlap between the two species, and therefore are not considered diagnostic.Regarding the number of small posterior teeth on the dentary, we found the total range of 4���7 with the following distribution: 5���7 in specimens of rio Paran�� drainage (MCP 20339 and MCP 37737); 4���6 in specimens of rio Paraguai drainage (MCP 39750); 4���5 in specimens of SLP (MCP 18666); 4���7 in specimens of rio Igua��u drainage (MCP 26610),and 4���5 in specimens of rio S��o Francisco drainage (MCP 17155 and MCP 16997). The differences in height between the upper and lower portions of the body (considering a horizontal line that passes by the oral cleft, as per the authors) were variable in the material examined. For example, lot MCP 27862 (7, 70.0��� 107.0 mm) of the rio Tiet��, upper rio Paran��, in which specimens present weak dark spots on the scales that are almost imperceptible on the ventral part of the body, presents a pattern that according to Garutti & Britski (2000) is characteristic of the species A. lacustris and A. altiparanae. We have verified that the upper and lower portions of the body are approximately equal among all specimens, with a tendency for the upper part to be a bit larger. See also the syntypes of Tetragonopterurs lacustris (Figs. 1 C, D), in which the height of the upper portion is smaller or approximately equal to the ventral portion. Salgado (2013) indicates characters differentiating A. altiparanae and A. lacustris, which include shape of the distal margin of the anal fin, shape of the margin of the operculum, length of the second ramified ray of the anal fin, and number of cusps on the symphyseal tooth. None of these characters were diagnosed in this study. Tetragonopterus jacuhiensis was described by Cope (1894) based on various specimens collected in ���Rio Grande do Sul��� [locality restricted by Fowler (1906) to the rio Jacu��, Sistema da Laguna dos Patos]. In characterizing the species, Cope mentions diameter of orbit (4 times in head length) and number of anal fin rays (28) as main diagnostic characters. Examination of the holotype revealed the orbit diameter occupying 33.2 % of the head lenght (about 3 times in head length), a value within the limits of A. lacustris. The same occurred regarding the number of anal fin rays. Although Cope did not mention spots on the scales forming horizontal stripes, examination of the holotype (Fig. 9 A) and some paratypes showed the presence of spots with weak intensity. The extensive material examined from the type locality, however, confirmed the presence of this character in this species, and highlighted the variability detailed above (Fig. 9 B). In reference to the coloration of Astyanax lacustris, although the concentration of chromatophores occurs in the same position on the scales, the intensity of spots and number of stripes may vary independently from the size of each specimen, or from its area of occurrence. Two color patterns are observed: (1) overall body light, silver on the lateroventral portion, with small dark spots on the posterior portion of some scales faded, forming 2 to 3 horizontal stripes above the lateral line, and 2, maximum 3, stripes below the lateral line (Figs. 1 A; 6 B, and 9 B); and (2) body darker, especially in the lateroventral region, with conspicuous spots, small on the posterior portion of most scales, forming in conjunction 3 to 4 horizontal stripes above the lateral line and 4, rarely 5 stripes below the lateral line (Figs. 1 B, 6 C, and 8 B). Considering the results obtained in the present study and exposed above, and base, Published as part of De Lucena, Carlos Alberto S. & Soares, Helena Gouvea, 2016, Review of species of the Astyanax bimaculatus " caudal peduncle spot " subgroup sensu Garutti & Langeani (Characiformes, Characidae) from the rio La Plata and rio S��o Francisco drainages and coastal systems of southern Brazil and Uruguay, pp. 101-125 in Zootaxa 4072 (1) on pages 103-114, DOI: 10.11646/zootaxa.4072.1.5, http://zenodo.org/record/256461, {"references":["Lutken, C. F. (1875) Velha-Flodens Fiske: Et Bidrag til Brasiliens Ichthyologie [Peixes do rio das Velhas: uma contribuicao para a ictiologia do Brasil]. In: Alves, C. B. M. & Pompeu, P. S. (Org.), Peixes do Rio das Velhas: passado e presente. SEGRAC, Belo Horizonte, pp. 23 - 164.","Cope, E. D. (1894) On the fishes obtained by the Naturalist Expedition in Rio Grande do Sul. Proceedings of the American Philosophical Society, 33, 84 - 108.","Fowler, H. 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