Tetragonopterus akamai, Araujo & Lucinda, 2014

Araujo, Leonardo B. S. & Lucinda, Paulo H. F., 2014, A new species of the genus Tetragonopterus Cuvier, 1816 (Ostariophysi: Characiformes: Characidae) from the rio Tocantins drainage, Brazil, Neotropical Ichthyology 12 (2), pp. 309-315 : 310-314

publication ID

https://doi.org/ 10.1590/1982-0224-20130173

DOI

https://doi.org/10.5281/zenodo.4775859

persistent identifier

https://treatment.plazi.org/id/03B8480A-3F13-FFE6-2CF3-FF6D3D76FBE2

treatment provided by

Carolina

scientific name

Tetragonopterus akamai
status

sp. nov.

Tetragonopterus akamai , new species

Figs. 1-2 View Fig View Fig

Holotype. MCP 48033, 56.5 mm SL, female, Brazil, Tocantins, Peixe, rio Tocantins close to confluence with rio Santa Tereza, 11º47’27”S 48º37’2”W, 7 Oct 2002, staff of the Núcleo de Estudos Ambientais , Universidade Federal do Tocantins (Neamb-UFT). GoogleMaps

Paratypes. All collected by Neamb-UFT staff (ordered from North to South). Brazil. Tocantins . UNT 8210 View Materials , 1 View Materials , 61.2 mm SL, Palmas, praia do Prata , 10º13’29”S 48º22’15”W, 28 Nov 2002 GoogleMaps . UNT 2807 View Materials , 1 View Materials , 60.6 mm SL, Porto Nacional, rio Tocantins, praia Porto Real , approximately 10º43’S 48º24’W, 6 Jun 1996 GoogleMaps . UNT 5160 View Materials , 1 View Materials , 47.0 mm SL, same locality as UNT 2807, 6 Jul 1996 GoogleMaps . UNT 2797 View Materials , 1 View Materials c&s, 54.3 mm SL, Brejinho de Nazaré, lagoa Pedra do Santo , approximately 11º01’S 48º34’W, 24 Nov 1996 GoogleMaps . UNT 10268, 11 View Materials , 8.2-44.7 mm SL, same locality as UNT 2797, 12 Mar 1997 GoogleMaps . MCP 48034, 3 View Materials , 52.9-57.9 mm SL, and UNT 5176 View Materials , 7 View Materials (1 c&s), 51.1-59.2 mm SL, Brejinho de Nazaré, lagoa Capivara , approximately 11º01’S 48º35’W, 14 Feb 1996 GoogleMaps . UNT 2788 View Materials , 1 View Materials c&s, 54.4 mm SL, Brejinho de Nazaré, lagoa Capivara , approximately 11º01’S 48º35’W, 14 Feb 1995 GoogleMaps . UNT 3544 View Materials , 10 View Materials , 33.2-42.1 mm SL, Ipueiras , rio Tocantins, approximately 11º19’S 48º28’W, 20 Apr 2001 GoogleMaps . UNT 10263, 1 View Materials , 48.9 mm SL, same locality as UNT 3544, 8 Aug 2002 GoogleMaps . UNT 5175 View Materials , 5 View Materials , 35.1-37.1 mm SL, same locality as UNT 3544, 28 Nov 2001 GoogleMaps . UNT 8626 View Materials , 1 View Materials , 39.8 mm SL, same locality as UNT 3544, 28 Nov 2002 GoogleMaps . UNT 3548 View Materials , 1 View Materials , 54.0 mm SL, Peixe, lagoa Dionísio, fazenda Água Branca , 11º44’49”S 48º38’16”W, 18 Mar 2001 GoogleMaps . UNT 2787 View Materials , 1 View Materials , 47.1 mm SL, same locality as holotype, 3 Jun 2002 GoogleMaps . UNT 5174 View Materials , 1 View Materials , 55.6 mm SL, same locality as holotype, 9 Sep 2002 GoogleMaps . UNT 10264, 1 View Materials , 46.6 mm SL, collected with the holotype . UNT 2792 View Materials , 1 View Materials , 47.6 mm SL , MCP 48036, 4 View Materials , 44.9- 45.8 mm SL, and UNT 3521 View Materials , 4 View Materials , 38.1-42.1 mm SL, Peixe, rio Santa Tereza, fazenda Água Branca , 11º48’07”S 48º38’21”W, 10 Sep 2002 GoogleMaps . UNT 2800 View Materials , 2 View Materials , 49.7-50.4 mm SL, Peixe, fazenda Água Branca, lagoa Água Branca , 11º49’20”S 48º38’40”W, 13 Oct 2000 GoogleMaps . UNT 5162 View Materials , 1 View Materials c&s, 47.1 mm SL, same locality as UNT 2800, 3 Jun 2002 . UNT 10265, 1 View Materials , 52.9 mm SL, same locality as UNT 2800, 22 May 2001 . UNT 10267, 1 View Materials , 37.0 mm SL, Peixe, rio Tocantins, UHE Peixe Angical , approximately 12º15’S 48º23’W, 24 May 2005 GoogleMaps . UNT 464 View Materials , 1 View Materials c&s, 46.5 mm SL, Paranã, rio Palmeiras drainage, 28 Oct 2002 . UNT 2785 View Materials , 1 View Materials , 52.8 mm SL, Paranã, rio Tocantins, fazenda Traçadal , approximately 12º29’S 48º12’W GoogleMaps . UNT 2793 View Materials , 1 View Materials c&s, 53.9 mm SL, Paranã, lagoa Traçadal, fazenda Traçadal , approximately 12º29’S 48º12’W GoogleMaps . UNT 10266, 1 View Materials , 55.6 mm SL, Paranã, rio Paranã, fazenda Traçadal , approximately 12º30’S 48º12’W, 24 Mar 1998 GoogleMaps .

Diagnosis. Tetragonopterus akamai is distinguished from all its congeners by a very uniquely shaped maxilla in the manner described below and depicted in Fig. 2 View Fig . The anterodorsal portion of the maxilla that is articulated with the premaxilla is bent over and in close contact with the dorsal surface of the premaxilla (vs. anterodorsal portion of the maxilla articulated with the premaxilla not bent over and not in contact with the dorsal surface of the premaxilla; see Fig. 2 View Fig and Silva & Benine, 2011: fig. 2). The anterodorsal portion of the maxilla articulated with the premaxilla is very short, ca. 5.5-5.9% of length of the free portion of the maxilla (vs. much longer, ca. 16.6-20% in the remaining species; see Fig. 2 View Fig and Silva & Benine, 2011: fig. 2). The anterodorsal border of the proximal portion of the maxilla articulated with the premaxilla is concave and angled approximately 90 degrees relative to the free portion of the maxilla (vs. continuous and not angled or angled approximately 45 degrees relative to the free portion of the maxilla in the remaining species; see Silva & Benine, 2011: fig. 2). The tip of the anterodorsal portion of the maxilla that is articulated with the premaxilla does not reach the vertical through the anterior margin of the antepenultimate teeth of the inner premaxillary series (vs. surpassing such vertical in the remaining species). Tetragonopterus akamai further differs from T. argenteus Cuvier ; T. chalceus Spix & Agassiz ; T. denticulatus Silva, Melo, Oliveira & Benine ; T. araguaiensis Silva, Melo, Oliveira & Benine , T. rarus Zarske, Géry & Isbrücker ; and T. carvalhoi Melo, Benine, Mariguela & Oliveira in the position of the mouth (superior vs. terminal, respectively). Tetragonopterus akamai can also be distinguished from T. anostomus Silva & Benine by the longer snout (20.0-27.6% vs. 9.5-14.3% HL, respectively) and by the relative size of the teeth on the inner premaxillary series, which are much larger than those of the outer series (vs. teeth of the inner premaxillary series as large as or smaller than those of the outer series in T. anostomus ). Tetragonopterus akamai can be further differentiated from T. anostomus by the number of gill rakers on the first branchial arch (9-10/14-16 vs. 11/18, respectively). Tetragonopterus akamai can be distinguished from T. rarus in the absence of dark longitudinal stripes along the flanks (vs. presence of such stripes in T. rarus ). Tetragonopterus akamai can be further differentiated from T. carvalhoi in the shape of the patch of dark pigmentation on the caudal peduncle (a round blotch restricted to the posterior half of the caudal peduncle vs. a lozenge-shaped blotch stretching out the whole extension of the caudal peduncle, respectively) (compare Fig. 1 View Fig with Melo et al., 2011: Fig 2a View Fig ), and in the higher number of gill rakers on the lower limb of the first branchial arch (14-16 vs. 13, respectively). Tetragonopterus akamai differs from T. argenteus in the number of predorsal scales (7-10 vs. 13- 17, respectively). Tetragonopterus akamai can be further distinguished from T. araguaiensis by: (1) the number of large teeth on the dentary (five or six vs. four, respectively); (2) the possession of overall relatively smaller dentary teeth with pointed cusps (vs. overall robust dentary teeth with roundish cusps, respectively); (3) the longer snout (20.0 - 27.6% vs. 14.3-18.1% HL, respectively); and (4) the number of gill rakers on the lower limb of the first gill arch (14-16 vs. 19-20, respectively). Tetragonopterus akamai can be further distinguished from T. denticulatus by (1) the distance between the first and second humeral blotches (distant 1-1.5 scales from each other in horizontal direction vs. distant 3 scales from each other, respectively); and (2) by the longer snout (20.0 - 27.6% vs. 10.9-18.4% HL, respectively).

Description. Morphometric data in Table 1 View Table 1 . Body moderately deep and laterally compressed. Greatest depth at dorsal-fin origin. Predorsal profile slightly straight from snout tip to extremity of supraoccipital bone and concave from that point to dorsal-fin origin. Postdorsal profile straight from base of last dorsal-fin ray to origin of adipose fin and convex from latter point to end of caudal peduncle. Ventral profile concave from snout to origin of pelvic fin. Ventral area between pelvic-fin insertion and anal-fin origin pronouncedly flattened. Sides of flattened area keeled. Margin of anal fin oblique along base of anal-fin rays. Ventral profile concave from base of last anal-fin ray to end of caudal peduncle.

Mouth superior. Two rows of premaxillary teeth. Teeth of internal row approximately twice as wide as teeth of external row. Teeth of external row of premaxilla approximately of same size. Five (3) or 6 (3) tri- or tetracuspid teeth in external row of premaxilla with pointed cusps of approximately similar size. First, or first and second anteriormost teeth as well as posteriormost teeth of external premaxillary row with three cusps and remaining teeth with four cusps. Teeth of internal row of premaxilla of approximately similar size. Five (5) or 6 (1) tri- or pentacuspid teeth in internal row of premaxilla with pointed cusps. Central cusps larger than lateral ones. First, or first and second anteriormost teeth as well as penultimate and last posteriormost teeth of internal premaxillary row with three cusps. Remaining teeth with five cusps. Anterodorsal portion of maxilla that is articulated with premaxilla very short, ca. 5.5-5.9% of maxillary free portion length. Anterodorsal border of proximal portion of maxilla articulated with premaxilla concave ( Fig. 2 View Fig ). Anterodorsal portion of maxilla articulated with premaxilla approximately angled 90 degrees relative to free portion of maxilla ( Fig. 2 View Fig ). Tip of anterodorsal portion of maxilla articulated with premaxilla not reaching vertical passing through anterior margin of antepenultimate teeth of inner premaxillary series. Three (4) or 4 (2) teeth of approximately similar size on proximal portion of maxilla. First, second, and third maxillary teeth unicuspid. Fourth tooth, when present, with central cusp and two or three lateral minute cusps ( Fig. 2 View Fig ). Five (3) or 6 (3) anteriormost dentary teeth larger, followed by 9 (1) or 12 (5) smaller teeth. Anterior large teeth of dentary with 3, 4, or 5 cusps. Posterior small teeth of dentary with one or two cusps.

Scales cycloid. Lateral line complete and distinctly ventrally curved anteriorly. Perforated scales on lateral line 29 (1), 30 (16), 31* (7), 32 (16), 33 (5), or 35 (1). Scale rows between dorsal-fin origin and lateral line 6* (23) or 7 (23). Scale rows between lateral line and pelvic-fin origin 3 (2), 4* (16), 5 (26), or 6 (2). Scale rows between lateral line and anal-fin origin 3* (28) or 4 (18). Longitudinal scale rows around caudal peduncle 10 (1), 11 (15), 12* (22), or 13 (8). Predorsal scales 7 (2), 8 (24), 9* (17), or 10 (3) arranged in regular series.

Dorsal-fin rays ii,8 (7) or ii,9* (39). Pectoral-fin rays i,11 (12), i,12* (21), or i,13 (9). Pelvic-fin rays i,6 (1) or i,7* (45). Anal-fin rays iii,27* (3), iii,28 (3), iii,29 (4), iii,30 (3), iii,31 (1), iii,32 (1), iii,33 (2), iii,34 (1), iv,26 (1), iv,27 (3), iv,28 (8), iv,29 (13), or iv,30 (3). Some specimens exhibiting sheath of small scales arranged in two rows along anal-fin base. Branched caudal-fin rays 18* (14), 19 (26), or 20 (6). Caudal fin scaled on some specimens, with first fifth basal portion of upper and lower lobe covered with small scales.

Precaudal vertebrae 11 (1), 12 (2), or 13 (3). Caudal vertebrae 18 (2), 19 (1), or 20 (3). Total vertebrae 30 (1), 31 (1), or 33 (3). Supraneurals 3 (4) or 4 (2). Dorsal procurrent rays 7 (2) or 10 (4). Ventral procurrent rays 5 (1), 6 (3) or 7 (2).

Color in alcohol. Background color yellow. Two pale brown humeral spots, vertically elongate and separated by one to one and half scales from each other. Each humeral spot covering three to four scales in vertical direction and one and half scale in horizontal direction. First humeral spot more prominent than second and distant one to two scales length from posteriormost point of operculum. Silver band extending along midline of flanks from region just posterior to distal border of opercle to base of caudal-fin rays. Rounded and well-defined light brown blotch on caudal peduncle. Fins hyaline.

Sexual dimorphism. Mature males can be recognized by the presence of bony hooks on the pelvic- and anal-fin rays. Hooks are present along extension of last unbranched anal-fin ray and along extension of first to seventh branched anal-fin rays and along extension of first five or first six branched pelvic-fin rays. The smallest mature male examined is 46.5 mm SL.

Distribution. Tetragonopterus akamai is known from the upper and middle portions of the rio Tocantins drainage ( Fig. 3 View Fig ).

Etymology. The specific name akamai (noun masculine genitive singular) is a patronym for Alberto Akama, in recognition of his contributions to Neotropical ichthyology and in recognition of his many collecting efforts on several South American drainages.

in the map. Discussion

Eigenmann (1917: 54) diagnosed Tetragonopterus in the following manner, which we below reproduce verbatim omitting some fragments and stressing the most important traits: “Small fishes, […] much compressed and very deep, the depth at least half the length; […] snout very short, the maxillary nearly vertical; […] premaxillary teeth in two rows […] several large, graduated, several pointed teeth in the front of the lower jaw, abruptly minute teeth on the sides; maxillary with or without teeth on its upper anterior edge […] the anal long, 32-37; scales entire, large on the middle of the sides, becoming smaller in all directions, notably toward front of anal […] lateral line complete, much decurved, several scales between its origin and that of the regular series below it […] preventral area flat, bounded by sharp angles, a median series of scales on the breast; […] This genus, most nearly allied to Moenkhausia , is readily distinguished from it by the greatly decurved lateral line which is not parallel with the row of scales below it in front”.

Given that a cladistic diagnosis (i.e., hypothesis of monophyly) of Tetragonopterus is not available to date, the traditional non-cladistic diagnosis of Eigenmann reproduced above is still broadly used. Therefore, in the absence of a cladistic diagnosis we opted to describe Tetragonopterus akamai as a species of Tetragonopterus species because it better conforms to the current non-cladistic Eigenmann’s (1917: 54) diagnosis of the genus. Moreover, Tetragonopterus akamai exhibits three supraneurals and a branched laterosensory canal in the sixth infraorbital, both conditions proposed as diagnostic for Tetragonopterus by Melo et al. (2011: 53).

Mirande (2010: 503) hypothesized eight character states as autapormorphic for Tetragonopterus argenteus . These character states were herein evaluated for T. akamai . 1) The laterosensory canal in the sixth infraorbital of T. akamai is tripartite, exhibiting an anterior dorsal portion running towards the neurocranium. 2) Tetragonopterus akamai exhibits of a lateral line tube lying onto the membrane between the middle caudal fin-rays. 3) The lamellar portion of the maxilla of T. akamai possesses a bifurcated canal with one elongate branch parallel to the dorsal margin of the maxilla plus a short branch (one fourth to one third of the length of the elongate branch) directed and parallel to the anterior margin of maxilla along one third of its length. 4) The ethmopalatine cartilage of T. akamai is large. 5) The base of the second pectoral fin-ray in T. akamai is swelled and extends laterally and beyond the base of first pectoral fin-ray. 6) The anterior border of scales of T. akamai is conspicuously and regularly undulated, and their radii converge to the center. 7) Finally, Tetragonopterus akamai exhibits two conspicuous and vertical humeral spots. 8) All the above-mentioned character states displayed by T. akamai are congruent with and can be coded in the same manner as those of T. argenteus .

We also evaluated the conditions of those character states ( Mirande’s (2010) characters 76, 93, 98, 171, 231, 318, 322, and 342) for the seven remaining species of Tetragonopterus . The character states exhibited by the specimens examined endorsed the conclusion that the following seven out of the eight autapomorphies listed by Mirande (2010) for T. argenteus are indeed apomorphies shared with all its congeners and therefore are herein interpreted as putatively synapomorphic for the genus Tetragonopterus , namely: (1) laterosensory canal of sixth infraorbital branched (character state 76-1), (2) caudal-fin canal of lateral line almost reaching posterior margin of caudal fin (character state 93-1); (3) tubules for passage of blood vessels on lamellar portion of maxilla with anterior branch running parallel to anterior margin of maxilla and reaching one third of its length (character state 98-1); (4) conspicuous ethmopalatine cartilage (character state 171-1); (5) base of second pectoral ray large and partially overlapping base of first pectoral ray from medial view (character state 231- 0); (6) anterior margin of scales with conspicuous undulations (character state 318-1); and (7) radii of scales converging at focus (character state 322-1). However, a conspicuous and vertically elongate second humeral spot (character state 342-1) is present in T. akamai , T. araguaiensis , T. argenteus , T. carvalhoi , T. chalceus , and T. denticulatus , whereas T. anostomus and T. rarus display the plesiomorphic state (absent or inconspicuous). Therefore, character state 342-1 is not herein proposed as a putative synapomorphy of the genus Tetragonopterus , but for a less inclusive clade.

Moreover, Melo et al. (2011: 53) postulated that the presence of three supraneural bones could be a diagnostic feature of Tetragonopterus . The new species T. akamai and all seven congeners have three supraneurals.

Testing the hypothesis of monophyly is not the aim of the present paper. Nonetheless we suggest a provisional diagnosis of Tetragonopterus relying on the combination of character states by Eigenmann (1917), on the presence of three supraneurals, and on the seven supposed synapomophies above-mentioned until the hypothesis of monophyly of the genus Tetragonopterus is tested.

Four additional nominal species of Tetragonopterus have been described based on specimens from the rios Tocantins- Araguaia drainage, namely T. anostomus , T. araguaiensis , T. denticulatus , and T. sawa Castelnau (= T. argenteus ). Besides T. argenteus and T. chalceus have also been reported from the rio Tocantins drainage. It is noteworthy that T. anostomus , T. araguaiensis , and T. denticulatus are known from the rio Araguaia and have not been recorded to date from the remaining portions of the rio Tocantins drainage. Nonetheless, T. anostomus , T. araguaiensis , T. argenteus , T. chalceus , and T. denticulatus clearly differ from T. akamai as detailed in the “Diagnosis” section.

A fourth species-name is available for the rio Tocantins drainage. Tetragonopterus sawa was described by Castelnau (1855: 65; pl. 33, fig. 1) from specimens of rio Crixás, an affluent to rio Araguaia. Hitherto the nomenclatural status of T. sawa is uncertain and it was treated as Species Inquirenda by Lima et al. (2003: 161). However, we examined a photograph of the holotype of T. sawa (MNHN A-9819) and we concluded that the specimen exhibits 14 pre-dorsal scales (vs. 7-10 in T. akamai ), which allowed us to infer that T. sawa and T. akamai are not conspecific. Tetragonopterus sawa is most likely a junior synonym of T. argenteus as previously recognized by Eigenmann (1917: 56) and suggested by Silva & Benine (2011: 54).

Comparative material. Tetragonopterus araguaiensis . Brazil. Goiás. LBP 4154, 1, 62.3 mm SL, paratype, Aragarças, rio Araguaia. LBP 7756, 1 c&s, 33.1 mm SL, paratype, Cocalinho , rio Araguaia. MCP 41394, 1 View Materials , 57.2 mm SL, Piranhas, rio Piranhas , near mouth of rio São Domingos , affluent of rio Araguaia. NUP 14678, 3, 53.1- 55.9 mm SL, São Miguel do Araguaia , lago Luís Alves, affluent to rio Araguaia. Mato Grosso. DZSJRP 3897, 4, 47.0- 57.7 mm SL, Santa Teresinha, rio Araguaia. Tetragonopterus argenteus . Brazil. Goiás. UNT 724 View Materials , 1 View Materials , 95.4 mm SL, Minaçu, rio Maranhão. UNT 725 View Materials , 1 View Materials , 85.3 mm SL, Niquelândia, rio Maranhão. UNT 726 View Materials , 1 View Materials , 99.4 mm SL, Niquelândia , rio Maranhão. Maranhão. UNT 2803 View Materials , 1 View Materials , 52.1 mm SL, Estreito , rio Tocantins close to confluence with rio Itaueiras. Tocantins. UNT 435 View Materials , 1 View Materials , 99.1 mm SL, Paranã, rio Palmeiras. UNT 2758 View Materials , 3 View Materials , 66.1-80.7 mm SL; UNT 2759 View Materials , 2 View Materials , 67.1 View Materials and 75.3 mm SL; UNT 2765 View Materials , 1 View Materials , 64.9 mm SL; UNT 2780 View Materials , 1 View Materials , 76.4 mm SL; UNT 2791 View Materials , 1 View Materials , 71.7 mm SL; UNT 3506 View Materials , 1 View Materials c&s, 76.8 mm SL; and UNT 3507 View Materials , 2 View Materials , 55.7 View Materials and 78.7 mm SL; Porto Nacional , rio Tocantins. UNT 2760 View Materials , 1, 108.2 mm SL, Paranã, rio Maranhão, fazenda Traçadal. UNT 2761 View Materials , 1 View Materials , 82.4 mm SL; UNT 2764 View Materials , 1 View Materials , 73.7 mm SL; and UNT 2770 View Materials , 1 View Materials c&s, 62.3 mm SL; Pedro Afonso , rio Tocantins close to confluence with rio Sono. UNT 2768 View Materials , 2 View Materials , 63.4 View Materials and 84.8 mm SL; UNT 2777 View Materials , 1 View Materials , 80.5 mm SL; and UNT 2781 View Materials , 1 View Materials , 67.4 mm SL; Brejinho de Nazaré , lagoa Capivara. UNT 2769 View Materials , 1 View Materials , 81.8 mm SL, Santa Rosa , rio Manoel Alves. UNT 2801 View Materials , 3 View Materials , 27.0- 31.2 mm SL, Brejinho de Nazaré , lagoa Pedra do Santo. UNT 3504 View Materials , 1 View Materials , 89.9 mm SL, Lajeado , rio Tocantins. UNT 4227 View Materials , 1 View Materials , 46.7 mm SL; and UNT 5167 View Materials , 1 View Materials , 35.8 mm SL; Ipueiras , rio Tocantins close to confluence with rio Manoel Alves. UNT 7173 View Materials , 3 View Materials , 62.2-68.7 mm SL, Peixe , rio Tocantins, upstream UHE Peixe Angical. UNT 7693 View Materials , 2 View Materials , 81.1 View Materials and 82.2 mm SL; and UNT 8163 View Materials , 1 View Materials , 90.7 mm SL; Paranã, rio Paranã, close to confluence with rio Lages. UNT 8055 View Materials , 1 View Materials , 80.5 mm SL, São Salvador , rio Maranhão, close to Distrito Retiro. UNT 10220, 1 View Materials , 52.6 mm SL, border of municípios de Sucupira and Peixe, rio Santa Tereza . Paraguay. MCP 15619, 6 View Materials , rio Paraguai, Cáceres. Tetragonopterus anostomus . Brazil. Goiás. LBP 7687, 2, 37.9 and 39.6 mm SL, paratype, Cocalinho, marginal lake near rio Araguaia. MCP 44638, 1 View Materials , 57.8 mm SL, São Miguel do Araguaia , rio Verde. MCP 44661, 6 View Materials , 34.4-40.9 mm SL, Britânia, stream of rio Araguaia drainage. MZUSP 108957 View Materials , 1 View Materials , 45.4 mm SL, holotype, Nova Crixás, rio Preto. MZUSP 89295 View Materials , 5 View Materials , 50.9 View Materials - 42.7 mm SL, paratopotypes. UFRGS 11627 View Materials , 4 View Materials , 30.4-41.1 mm SL, Barra do Garças , ribeirão Corrente near road BR-158 towards Nova Xavantina. Tetragonopterus carvalhoi : Brazil. Amapá. LBP 5376, 2, 42.1 and 44.2 mm SL, paratypes, collected with the holotype. MZUSP 101755 View Materials , 2 View Materials , 39.5 View Materials and 4.2 mm SL, rio Jari, upstream and downstream cachoeira Santo Antônio. MZUSP 102268 View Materials , 1 View Materials , 36.5 mm SL, holotype, Laranjal do Jari, igarapé Iratapuru , rio Jari. MZUSP 103376 View Materials , 4 View Materials , 44.8-64.5 mm SL, Laranjal do Jari , igarapé três bocas, affluent to rio Pacanari downstream cachoeira Santo Antônio. MZUSP 106813 View Materials , 2 View Materials , 59.2 View Materials and 63.4 mm SL, paratopotypes. MZUSP 101708 View Materials , 1 View Materials , 60.7 mm SL, Monte Dourado, rio Jari. Tetragonopterus chalceus . Brazil. Tocantins. UNT 2762 View Materials , 1 View Materials , 70.8 mm SL; UNT 2766 View Materials , 3 View Materials , 2 View Materials c&s, 51.8-55.6 mm SL; and UNT 2794 View Materials , 1 View Materials , 59.2 mm SL, Paranã, rio Paranã, fazenda Traçadal. UNT 2771 View Materials , 2 View Materials , 1 View Materials c&s, 57.1 and 74.0 mm SL; UNT 2778 View Materials , 1 View Materials , 75.0 mm SL; and UNT 10277, 2 View Materials , 51.3 View Materials and 62.6 mm SL; Paranã, rio Maranhão, fazenda Traçadal. UNT 2772 View Materials , 2 View Materials , 71.5-74.3 mm SL; and UNT 2783 View Materials , 2 View Materials , 1 View Materials c&s, 54.8-65.0 mm SL; Brejinho de Nazaré, lagoa Capivara. UNT 2775 View Materials , 1 View Materials , 90.4 mm SL; UNT 2790 View Materials , 1 View Materials , 54.0 mm SL; and UNT 3523 View Materials , 4 View Materials , 50.6-66.4 mm SL, Peixe, rio Santa Tereza. UNT 2779 View Materials , 3 View Materials , 55.4-57.5 mm SL; UNT 6476 View Materials , 1 View Materials , 37.1 mm SL; UNT 10274, 1 View Materials , 56.6 mm SL; and UNT 10275, 2 View Materials , 53.9-59.1 mm SL; Porto Nacional, rio Tocantins. UNT 2784 View Materials , 1 View Materials , 54.6 mm SL, Porto Nacional, rio Mangues. UNT 2795 View Materials , 1 View Materials , 53.7 mm SL, Ipueiras, rio Formiga close to confluence with rio Tocantins. UNT 2799 View Materials , 1 View Materials , 57.1 mm SL, Brejinho de Nazaré , rio Crixás. UNT 3505 View Materials , 1 View Materials , 79.3 mm SL, Peixe, lagoa Dionísio, fazenda Água Branca. UNT 3508 View Materials , 3 View Materials , 62.5-79.3 mm SL, Tupirama, rio Tocantins close to confluence with rio Tranqueira. UNT 3509 View Materials , 3 View Materials , 52.9-55.9 mm SL, Peixe, lagoa Água Branca, fazenda Água Branca. UNT 3510 View Materials , 2 View Materials , 55.1-71.7 mm SL, Peixe, rio Tocantins close to confluence with rio Santa Tereza. UNT 3511 View Materials , 1 View Materials , 66.2 mm SL, Lajeado, rio Tocantins, fish ladder. UNT 3540 View Materials , 2 View Materials , 49.2 View Materials and 51.3 mm SL, Porto Nacional, rio Tocantins near to the confluence with rio Feio. UNT 3543 View Materials , 6 View Materials , 50.1-56.4 mm SL, Paranã, rio Paranã close to lagoa Verde. UNT 3546 View Materials , 10 View Materials , 36.5-44.3 mm SL, Peixe, rio Tocantins close to confluence with rio Almas. UNT 5161 View Materials , 2 View Materials , 50.7-55.4 mm SL, Paranã, rio Paranã, lagoa Traçadal. UNT 8052 View Materials , 1 View Materials , 1 View Materials , 71.2 mm SL, Paranã, córrego das Pedras. UNT 10276, 1 View Materials , 30.3 mm SL, Peixe, rio Tocantins close to UHE Peixe Angical. Tetragonopterus denticulatus . Brazil. Goiás. DZSJRP 13587, 5, 46.7-52.1 mm SL, Aragarças, Praia Quarto Crescente, rio Araguaia. LBP 5751, 2, 1 c&s, 50.4-55.8 mm SL, parátipos, Aragarças , rioAraguaia. Mato Grosso. MZUSP 42782 View Materials , 4 View Materials , 29.6-37.8 mm SL, Barra do Garças , rio Araguaia. MZUSP 89182 View Materials , 4 View Materials , 41.0- 47.6 mm SL, Cocalinho, rioAraguaia. Tetragonopterus rarus . Brazil. Pará. MZUSP 88029 View Materials , 1 View Materials , 74.0 mm SL, Alto Pará do Oeste, rio Amazonas, affluent to rio Tírios . Suriname. LIRP 4928 View Materials , 2 View Materials cs, 47.5- 47.9 mm SL, and LIRP 4929 View Materials , 1 View Materials , 79.8 mm SL, tributary to Sisa Creek , Nickerie. MZUSP 99689 View Materials , 1 View Materials , 86.2 mm SL, Maroni, Marowjine Rivier .

MCP

Pontificia Universidade Catolica do Rio Grande do Sul

UNT

Universidad nacional de Tucumn

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