Cuvier, 1816 : 166 Müller & Troschel, 1845 : 1 Günther,1864 : 318 Eigenmann, 1917 : 54 Fowler, 1932 : 352 Géry, 1964a : 5 Ringuelet et al. , 1967 : 131 Ortega & Vari, 1986 : 9 Stewart et al. , 1987 : 29 Taphorn, 1992 : 343 Britski et al. , 1999 : 29 Tetragonopterinae Castelnau, 1855 : 65 Steindachner, 1909 : 172 Revision of the South American genus Tetragonopterus Cuvier, 1816 (Teleostei: Characidae) with description of four new species Silva, Gabriel S. C. Melo, Bruno F. Oliveira, Claudio Benine, Ricardo C. Zootaxa 2016 4200 1 1 46 Cuvier, 1816 Cuvier 1816 [151,638,546,572] Insecta Anthribidae Tetragonopterus Animalia Coleoptera 10 11 Arthropoda species argenteus     Tetragonopterus argenteus  Cuvier, 1816: 166(description; type locality: “South America);  Müller & Troschel, 1845: 1–2, 13 (description; Brazil, Guyana);  Günther,1864: 318(description; Bahia?);  Eigenmann, 1917: 54–55 (description; from Orinoco and Guiana to the Paranahyba basin, Amazons and Paraguay, Paraná to Buenos Aires);  Fowler, 1932: 352(description; illustration; upper Paraguay, Descalvados, MT, Brazil); 1940: 90 (citation; Boca Chapare, Bolivia); 1941: 185 (citation; northeastern Brazil, Rio Paranaíba, Piau; Rio Jaguaribe, Cear);  Géry, 1964a: 5(citation; Peruvian Amazonia);  Ringuelet et al., 1967: 131–132 (description; upper Paran, Candelária and Arroio Uruguai, Missiones; Middle Paran, Empredado; Ilha Apipé Grande, Corrientes; Río La Plata; upper and middle río Paraguay; Goiás; Amazonas; Guyana);  Ortega & Vari, 1986: 9(citation: Amazonas, Peru);  Stewart et al., 1987: 29(citation, río Napo, Ecuador Oriental);  Taphorn, 1992: 343(description; Apure River drainage, illustration);  Britski et al., 1999: 29–31 (description; Pantanal matogrossense); Reis, 2003: 212 (list of Tetragonopterinaespecies); Lima et al., 2013; 352–353 (description; Rio Madeira basin, Brazil; photo).    Tetragonopterus sawa  Castelnau, 1855: 65, pl. 33 (original description; type locality: “rio Crixas qui se jette dans L’Araguaia [Brazil]).    Tetragonopterus huberi  Steindachner, 1909: 172(original description; type locality: “Purus, Amazonas).   Diagnosis.  Tetragonopteurs argenteusis distinguished from all other congeners by having 11–17 predorsal scales ( vs.7–9). It further differs from  T. anostomus,  T. denticulatus,  T. kuluene, and  T. juruenaby having 4 principal teeth in dentary ( vs.5–6), and larger and more robust teeth ( vs.thinner and sharper teeth; Silva et al.2013: Fig. 2).  Tetragonopterus argenteusdiffers from  T. anostomusand  T. araguaiensisby having 12–15 gill-rakers on the upper limb of the first gill arch ( vs.17–20; Silva et al.2013: Fig. 3); it also differs from  T. anostomusby having a terminal mouth ( vs.subsuperior mouth).  Tetragonopterus argenteusdiffers from  T. ommatus sp. n.by bearing 1–4 maxillary teeth ( vs.7–8) and by the presence of a conspicuous dark mark on the caudal peduncle ( vs.inconspicuous mark limited to the posterior portion of the caudal peduncle).  Tetragonopterus argenteusdiffers from  T. carvalhoiby the presence of a rounded dark mark on the caudal peduncle ( vs.lozenge-shaped dark mark on the caudal peduncle).  Tetragonopterus argenteusdiffers from  T. rarusby the absence of dark longitudinal stripes on the trunk ( vs.presence of such stripes); it differs from  T. rarusand  T. georgiae n. comb.by having 3.5 scale rows between lateral line and pelvic-fin origin ( vs.4.5–5.5). Finally,  T. argenteusdiffers from  T. denticulatusby having humeral marks separated by one or two vertical rows of scales ( vs.humeral marks separated by three vertical scale rows).   Description.Morphometric data presented in Table 2. Body deep, compressed laterally, with greatest depth at dorsal-fin origin. Dorsal profile slightly convex between snout tip and vertical through middle of orbit; slightly concave from this point to end of occipital process; convex from that point to dorsal-fin origin. Slightly convex from dorsal-fin origin to adipose-fin end. Prepelvic region transversely flattened with distinct, longitudinally aligned lateral keels. Ventral profile of body convex from lower lip to anal-fin origin; straight or somewhat convex along base of anal fin. Caudal peduncle with dorsal and ventral profiles slightly concaves. Olfactory lamellae 17 (1), 18 (4), 19 (7), 20 (5), 21 (7), 22 (16), 23 (8), 24 (6), 25 (4), 26 (7), 28 (2), 32 (1), 35 (2) or 40 (1). Snout length smaller than orbital diameter. Mouth terminal. Premaxilla in two rows of relatively robust teeth. Outer row with 4 (6), 5 (68), 6 (104), 7 (31) or 8 (10) teeth with three cusps, central cusps longest. Inner row with 4 (1), 5 (228) or 6 (3) teeth with three, four or five cusps; if present, central cusp twice as long and broader than other cusps. Maxilla with 1 (3), 2 (62), 3 (132) or 4 (29) teeth with three cusps. Dentary with 3 (1) or 4 (228) anteriormost teeth with three or five cusps followed by a smaller tricuspid tooth and 8–13 smaller conical teeth of similar size (5 c&s).   TABLE 2.Morphometric đata for  Tetragonopterus argenteusfrom đistinct South American basins separateđ accorđing to the COI tree resolution (Fig. 1).    Measurements Stanđarđ length (mm) Amazon (n=117) Range Mean 40.8Q88.4 63.4 Tapajós anđ Xingu (n=5) Range Mean 64.8Q91.4 81.7 Araguaia (n=33) Range Mean 42.2Q88.2 53.4 La Plata basin (n=42) Range Mean 27.4Q110.5 58.0 Orinoco basin (n=4) Range Mean 57.1Q82.2 65.4 Parnaíba basin (n=28) Range Mean 30.1Q76.1 50.7   Percentages of standard lengthGreatest đepth 44.8Q64.2 55.7 52.2Q61.0 57.1 50.3Q62.6 56.1 45.8Q63.4 58.4 55.6Q57.9 56.6 45.8Q54.7 51.4  Pređorsal length 44.3Q61.0 52.9 52.1Q54.9 53.6 47.9Q55.7 52.9 45.3Q56.9 52.5 47.6Q50.7 48.6 49.2Q54.0 52.2  Prepectoral length 26.3Q34.2 29.8 28.0Q29.3 28.7 26.7Q34.1 30.9 26.9Q35.2 30.8 29.3Q31.5 30.3 25.9Q32.1 28.6  Prepelvic length Preanal length 46.5Q54.4 49.4 49.2Q77.8 66.6 46.8Q49.6 48.4 51.9Q67.0 63.7 46.5Q54.6 49.6 63.1Q71.1 66.0 46.4Q54.4 49.5 63.6Q73.8 67.1 49.2Q50.1 49.8 64.3Q69.2 66.6 46.0Q49.1 47.8 62.6Q68.0 64.8  Cauđal peđuncle đepth 8.4Q12.0 10.1 10.3Q11.9 11.0 9.0Q11.3 10.5 8.83Q13.8 10.9 6.8Q9.3 10.0 8.89Q11.6 10.2  Cauđal peđuncle length 6.0Q10.5 8.1 7.2Q7.9 7.6 6.3Q9.1 8.0 5.63Q9.67 7.78 6.0Q6.8 6.4 6.41Q9.96 8.29  Pectoral-fin length 21.1Q27.0 24.2 21.7Q25.6 24.1 20.3Q28.7 25.5 20.4Q32.9 25.1 20.4Q24.9 22.2 21.3Q24.3 22.7  Pelvic-fin length Dorsal-fin length 14.1Q23.5 20.6 15.7Q42.8 37.1 18.3Q22.3 20.3 35.3Q38.5 36.5 16.4Q25.1 19.8 31.7Q44.7 39.5 17.5Q25.2 20.4 34.9Q43.2 39.0 14.2Q18.8 16.4 39.1Q40.3 35.3 16.8Q22.8 19.5 33.3Q42.8 36.2  Length of đorsal-fin base 14.9Q38.5 17.1 16.5Q18.7 17.5 15.3Q18.8 17.5 14.1Q21.3 17.3 15.8Q17.1 16.3 14.0Q18.2 15.9  Anal-fin length 9.8Q23.6 15.5 9.2Q16.0 12.1 9.7Q19.9 16.1 8.6Q17.3 13.0 9.6Q10.4 9.98 14.0Q18.2 15.9  Length of anal-fin base Distance from eye to đorsal-fin origin 32.0Q51.3 39.2 44.1Q57.9 49.7 38.5Q43.2 41.2 49.4Q51.5 50.8 37.4Q44.7 39.7 44.6Q52.4 49.4 37.9Q48.7 41.7 47.5Q57.7 51.4 36.4Q37.6 37.2 50.2Q52.1 51.1 36.5Q42.3 39.3 46.1Q50.6 48.3  Distance from đorsal-fin origin to cauđal-fin base Heađ length 42.7Q65.9 55.1 25.6Q36.3 29.4 54.7Q61.8 56.9 27.4Q29.2 28.4 50.8Q58.4 54.7 26.1Q34.1 30.4 51.2Q67.1 55.4 24.9Q32.5 29.5 53.0Q55.1 54.4 25.7Q29.9 28.4 50.2Q57.9 54.5 25.5Q32.0 28.8  Heađ đepth 21.0Q28.2 23.9 22.7Q24.9 24.0 24.9Q29.3 27.0 23.9Q32.3 26.8 25.2Q27.1 26.3 19.9Q30.2 24.0   Percentages of head lengthSnout length 9Q18 13.8 15Q19 16.6 9Q19 14.8 11Q19 13.9 15Q16 15.6 13Q19 15.0  Maxillary length 31Q48 42.0 42Q45 43.0 31Q46 42.6 38Q46 42.3 43Q47 44.7 40Q45 42.2  Horizontal orbital điameter 38Q60 45.4 44Q48 45.8 42Q50 47.4 47Q58 51.4 43Q49 46.4 43Q51 46.8  Least interorbital wiđth 24Q41 34.8 36Q42 39.1 32Q40 35.3 30Q45 36.5 35Q41 37.4 31Q39 34.5   FIGURE 3.  Tetragonopterus argenteus.a) MNHN 9807, holotype, 112.0 mm SL, unknown locality; b) LBP 3758, 53.8 mm SL, Brazil, Mato Grosso, Paraguay basin; c) MUSM 39445, live specimen, 78.0 mm SL, Río Purus, Peru. Photos by MNHN (a) and T.P. Carvalho (c). Dorsal-fin rays ii,9 (232). First unbranched ray very short. Dorsal-fin origin anterior to middle of body in SL. Anal-fin rays v,28 (1) v,29 (1), v,30 (4), v,31 (9), iv,32 (36), iv,33 (43), iv,34 (56), iv,35 (44), iv,36 (22), iv,37 (9) or iv,40 (1); posterior unbranched rays and anterior branched rays usually slightly longer than following rays. Analfin origin at vertical through base of last branched dorsal-fin ray. Pelvic-fin rays i,7 (228). Pelvic-fin origin anterior to vertical through dorsal-fin origin. Distal margin of pelvic fin reaching pelvic-fin origin with anterior rays longest. Tip of adpressed pelvic fin surpassing anal-fin origin. Pectoral-fin rays i,11 (4), i,12 (59), i,13 (105), i,14 (49) or i,15 (7). Caudal fin forked with i,17,i principal rays (5 c&s). Dorsal procurrent rays 11–13 (5 c&s) and ventral procurrent rays 8–9 (5 c&s). Scales large and cycloid. Lateral line complete and distinctly curved downward anteriorly. Longitudinal scales 30 (5), 31 (17), 32 (48), 33 (79), 34 (52) or 35 (4). Scale rows between lateral line and dorsal-fin origin 7 (64) or 8 (162). Scale rows between lateral line and pelvic-fin origin 3.5 (234). Predorsal scales 11 (6), 12 (19), 13 (31), 14 (54), 15 (58), 16 (16), 17 (10) or 18 (2). Scale rows around caudal peduncle 13 (43) or 14 (138). Anal-fin base covered by a single row of small scales; anterior two thirds of lobes of caudal fin densely scaled in many specimens. First gill arch with 8 (5), 9 (80), 10 (71) or 11 (13) gill rakers on upper limb and 12 (18), 13 (68), 14 (69) or 15 (6) gill rakers on lower limb. Total vertebrae 27, precaudal vertebrae 11, caudal vertebrae 16 (5 c&s). Supraneurals 3 (5 c&s).  Color in alcohol.General body color yellowish. Dorsal portion of head and body darkly pigmented (OS 18365 from Río Itaya in Peruwith predorsal region densely pigmented). Dorsolateral portion of body with few minute chromatophores along distal margin of scales. Scales of ventrolateral portion of body unpigmented. Opercular and infraorbital bones silvery speckled with a few larger chromatophores. Two conspicuous vertical dark humeral marks separated by one or two vertical rows of scales. Anterior humeral mark extending over five horizontal scale rows above lateral line and two horizontal scale rows bellow lateral line. Posterior humeral mark extending over five or six horizontal scales rows above lateral line. Caudal peduncle with a rounded or vertically-oriented ellipsoid mark. Midlateral silvery stripe broad, extending from supracleithrum to anterior margin of the dark mark on the caudal peduncle. Anal, pelvic, and dorsal fins mostly hyaline, with chromatophores concentrated at distal portions of rays. Adipose fin speckled with small, dark chromatophores ( Fig. 3b).  Sexual dimorphism.Adult males with bony hooks on distal segments of fourth unbranched and anteriormost branched rays of anal fin; usually two bony hooks per segment. One adult male (ANSP 134765) from Río Meta in Colombia with bony hooks on fourth unbranched ray and on anteriormost 22 branched rays of anal fin.   Distribution.  Tetragonopterus argenteusis distributed across the Amazon basin, upper Essequibo, Orinoco, Parnaíba, Jaguaribe,  Paraguayand lower Paranáriver basins ( Fig. 2a).    Material examined. Non types: Ecuador:  ANSP75902, 1, 102.3 mm SL, Chicherota, Napobasin, Río Pastaza, 2°25’15.1”S 76°38’39.4”W.   ANSP137656, 9, 42.7–97.8 mmSL, Río Lagartococha.  Peru:  ANSP18365, 3, 75.1–82.7 mmSL,  Loreto, Maynas, Río Itaya, 4°13’31.2”S 73°28’57.3”W.  ANSP143548, 1, 54.9 mmSL, Sandoval, Cosha,  Río Madre de Diós. LBP 12401, 1, 65.6 mmSL,  Loreto, San Juan Bautista, Río Marañon, 4°09’04.2”S 73°28’25.7”W. LBP 12578, 1, 57.7 mmSL,  Loreto, Nauta, Río Marañon, 4°18’13.6”S 73°31’16.1”W.  MZUSP26393, 10, 40.8–67.5 mmSL, Pucallpa,  Río Ucayali.  MZUSP26434, 4, 55.7–61.1 mmSL, Cocha, Supay. OS18365, 3, 75.1–82.7 mmSL,  Loreto, Maynas, Río Itaya, 4°13’31.2”S 73°28’57.3”W.  Colombia:  ANSP131848, 4, 51.7–69.3 mmSL, Quebrada Venturosa, Río Meta.   ANSP134765, 3, 46.2–66.5 mmSL, Río Meta.   ANSP137585, 1, 54.9 mmSL, La Balsa, Río Meta, Río Negrito.   ANSP192316, 3, 45.0– 48.2 mmSL, Río Meta. LBP 18642, 2, 41.6–45.8 mmSL,  Granada, Orinoco basin, Río Guaviare.  Venezuela: LBP 3058, 2, 58.1–81.2 mmSL,  Caicara de Orinoco, Río Orinoco, 7°38’11”N 66°19’04.2”W. LBP 3059, 1, 57.1 mmSL, Caicara del Orinoco, Río Orinoco,  7°38’11”N 66°19’04.2”W. Guyana:  ANSP 190547, 6, 43.1–71.0 mm SL, Rupununi,  Essequibobasin, Rupununi river.  ANSP193496, 3, 55–60 mmSL, Rupununi, Essequibo basin.  Brazil: Acre: LBP 185, 2, 71.2–72.5 mmSL; LBP 16805, 8, 56.4–80.7 mmSL; LBP 16862, 1, 62.0 mm SL, Rio Acre-Purus, Rio Branco, 10°03.320’S 67°51.450’W.  MZUSP30363, 4, 41.7–47.4 mmSL, Tarauacá, Rio Juruá.   MZUSP50116, 2, 75.2–85.6 mmSL, Manuel Urbano, Rio Purus.   MZUSP50301, 3, 81.8–88.4 mmSL, Colocação Volta Grande, Rio Juruá.  Amazonas:  LIRP4973, 1, 50.9 mmSL,  Paranádo Janauacá.  MZUSP5773, 14, 66.3–88.4 mmSL,  Paranáde Urucará.  MZUSP6516, 5, 64.9–80.6 mmSL, Manacapuru, Rio Solimões,  3°13’00”S 60°45’00”W.  MZUSP17353, 9, 41.7–66.4 mmSL, Coari, Rio Solimões,  3°55’00”S 63°20’00”W.  MZUSP17509, 9, 61.7–72.1 mmSL, Santo Antonio do Içá, Rio Içá,  3°08’00”S67°57’00”.  MZUSP27756, 5, 70.9–82.5 mmSL, Benjamin Constant, Rio Içámouth, 24°04’00”S 69°49’00”W.   MZUSP74668, 3, 64.8–67.0 mm SL, Rio Solimões, 3°22’00”S 60°11’00”W.   MZUSP74687, 9, 55.4–68.1 mmSL, Rio Solimões. Pará: LBP 17608, 1, 65.0 mm SL, Altamira, Rio Xingu, 3°30’14.3”S 52°02’19.9”W.   MZUSP3528, 3, 87.6–91.4 mmSL, Santarém, Rio Tapajós, 2°25’00”S 54°44’00”W.   MZUSP5449, 4, 65.1–79.4 mmSL, Oriximiná, Rio Trombetas, 1°46’00”S 55°52’00”W. MZUSP 9494, 6, 48.6–55.3 mmSL, Monte Alegre, 2°01’00”S 54°55’00”W.  Piauí:  ANSP84415, 5, 53.1–61.9 mmSL, Terezina, Rio Parnaíba.   MZUSP5091, 15, 30.1–57.4 mmSL, Teresina, Rio Parnaíba, 5°05’00”S 42°50’00”W.   MZUSP74891, 14, 56.7–76.1 mmSL, Teresina, Rio Parnaíba.  Ceará:  ANSP84416, 2, 72.5–82.1 mmSL, Russas, Rio Jaguaribe, Russas.  Goiás: LBP 1832, 1, 68,8 mm SL, Aragarças, Rio Araguaia,  15°53’35.6”S 52o15’01”W. LBP 17275, 1, 60.9 mmSL, Britânia, Rio Araguaia, Rio Vermelho,  15°10’23.2”S 51°09’27.1”W.  MZUSP4814, 28, 42.6–58.5 mmSL, Aruanã, Rio Araguaia.   MZUSP54523, 1, 73.5 mmSL, Rio Araguaia.   MZUSP110039, 3, 76.4–88.2 mmSL, Minaçu, Rio Tocantins.  Maranhão: LBP 5535, 1, 74.4 mmSL, Balsas, rio Parnaíbabasin, Rio Balsas, 7°32’26”S 46°02’21”W.  Mato Grosso: LBP 3967, 1, 59.4 mmSL, Cuiabá, Rio Paraguai, Rio Mutuca,  15°21’20”S 56°06’04”W. LBP 4633, 3, 64.5–72.3 mmSL, Poconé, Rio Paraguai. LBP 5108, 1, 67.6 mmSL, Cáceres, Rio Paraguay,  16°06’66”S 57°44’33”W. LBP 8442, 3, 51.8–63.8 mmSL, Cáceres, Rio Paraguai,  16°03’13.6”S 57°48’31.8”W. LBP 14046, 1, 79.5 mmSL, Poconé, Rio Paraguai, Rio Cuiabá,  17°50’49.9”S 57°26’22.3”W. LBP 15921, 1, 78,6 mm SL, Canarana, Rio Xingu, Rio Culuene,  13°31’02.8”S 53°04’41.8”W. LBP 15962, 3, 52.4– 44.3 mmSL, Canarana, Rio Culuene,  13°29’41.8”S 53°04’57.7”W.  LIRP713, 2, 65.4–65.9 mmSL, Barão de Melgaço, Rio Paraguai, Rio Cuiabá,  16°12’00”S 55°57’30”W.  MZUSP 101060, 1, 52.1 mmSL, Rio Madeirabasin, Rio Aripuanã,  10°12’33”S 59°27’39”W.  MZUSP 101069, 15, 41.3–47.1 mmSL, Aripuanã, Rio Madeirabasin, Rio Aripuanã. NUP 3381, 5, 62.9–82.0 mm SL, Aripuanã, Rio Madeirabasin, Rio Aripuanã.NUP 4092, 3, 65.0– 87.6 mmSL, Chapada dos Guimarães, Rio Paraguai, Rio Cuiabá.NUP 7189, 2, 55.6–57.6 mmSL, Aripuanã. NUP 10758, 17, 21.9–44.4 mmSL, Rosário Oeste, Rio Paraguaibasin, Rio Cuiabá. NUP 10768, 1, 101 mmSL, Nova Brasilândia, Rio Paraguaybasin,  Rio Manso. Mato Grossodo Sul: LBP 1476, 7, 27.4–42.4 mmSL, Coxim, Rio Paraguai,  18°25’42.5”S 54°50’02”W. LBP 1776, 4, 81.3–97.8 mmSL, Coxim, Rio Paraguai. LBP 3758, 20, 50.4–66.6 mmSL, Aquidauana, Rio Paraguai, 19°34’33.7”W 56°14’49.5”S. LBP 13558, 1, 51.5 mmSL, Corumbá, Rio Paraguai, 17°55’16”S 57°28’31.6”W. LBP 14036, 2, 55.7–94.1 mmSL, Poconé, Río Paraguai, 17°30’49”S 57°14’26.3”W.   MZUSP59706, 21, 45.1–60.8 mmSL, Aquidauana, Rio Paraguaibasin, Rio Negro.  Paraná: NUP 7341, 1, 110.9 mmSL, Foz do Iguaçu, Rio Paraná. 1412767305 ANSP Chicherota Ecuador -2.4208612 Napo 1 -76.64428 13 14 ANSP 75902, 1 1 Napo holotype 1412767321 [573,1202,1124,1149] ANSP Ecuador Rio Lagartococha 13 14 ANSP 137656, 9 1 Napo holotype 1412767315 ANSP Peru Peru 13 14 ANSP 18365, 3 1 holotype 1412767306 ANSP Sandoval & Cosha Peru -4.225333 Rio Itaya 1 -73.48258 13 14 ANSP 143548, 1 1 Loreto holotype 1412767307 [362,941,1196,1221] Peru Rio Madre de Dios 13 14 1 Rio Madre de Dios holotype 1412767302 Peru -4.151167 Rio Maranon 1 -73.47381 13 14 1 Loreto holotype 1412767325 MZUSP Pucallpa Peru -4.303778 Rio Maranon 1 -73.52114 13 14 MZUSP 26393, 10 1 Loreto holotype 1412767323 MZUSP Cocha & Supay. OS Peru Rio Ucayali 13 14 MZUSP 26434, 4 1 Rio Ucayali holotype 1412767311 [562,1216,1303,1329] Peru -4.225333 Rio Itaya 1 -73.48258 Maynas 13 14 1 Loreto holotype 1412767308 ANSP Quebrada Venturosa Colombia Meta 13 14 ANSP 131848, 4 1 Meta holotype 1412767313 [882,1433,1340,1365] ANSP Colombia Meta 13 14 ANSP 134765, 3 1 Meta holotype 1412767301 [151,886,1375,1400] ANSP La Balsa Colombia Rio Negrito 13 14 ANSP 137585, 1 1 Meta holotype 1412767303 ANSP Colombia Meta 13 14 ANSP 192316, 3 1 Meta holotype 1412767310 [539,975,1412,1437] Grenada Guaviare 13 14 1 Guaviare holotype 1412767312 Caicara de Orinoco Venezuela Caicara de Orinoco 7.636389 Rio Orinoco 15 -66.31783 Rio Orinoco 13 14 1 holotype 1412767320 [303,1238,1484,1509] ANSP Rupununi Guyana 7.636389 Guyana 15 -66.31783 13 14 ANSP 190547, 6 1 holotype 1412767314 ANSP Rupununi Guyana Rupununi river 13 14 ANSP 193496, 3 1 holotype 1412767324 Brazil Acre 13 14 1 Acre holotype 1412767319 MZUSP Tarauaca Brazil Rio Acre-Purus -10.055333 Rio Jurua 1 -67.8575 Rio Branco 13 14 MZUSP 30363, 4 1 Acre holotype 1412767317 MZUSP Manuel Urbano Brazil Rio Purus 13 14 MZUSP 50116, 2 1 Acre holotype 1412767318 [468,1299,1628,1653] MZUSP Colocacao Volta Grande Brazil Rio Jurua 13 14 MZUSP 50301, 3 1 Acre holotype 1412767309 LIRP Brazil Amazonas 13 14 LIRP 4973, 1 1 Amazonas holotype 1412767340 [467,1112,1663,1688] MZUSP Brazil Parana 13 14 MZUSP 5773, 14 1 Parana holotype 1412767327 MZUSP Manacapuru Brazil Rio Solimoes 13 14 MZUSP 6516, 5 1 Parana holotype 1412767347 MZUSP Coari Brazil -3.2166667 Rio Solimoes 21 -60.75 13 14 MZUSP 17353, 9 1 Parana holotype 1412767334 MZUSP Brazil -3.9166667 Ica 21 -63.333332 13 14 MZUSP 17509, 9 1 Ica holotype 1412767344 [200,1292,1772,1797] MZUSP Benjamin Constant Brazil -3.1333334 Ica 13 14 MZUSP 27756, 5 1 Ica holotype 1412767342 [309,1208,1807,1832] MZUSP Brazil -3.3666668 Rio Solimoes 21 -60.183334 13 14 MZUSP 74668, 3 1 Ica holotype 1412767339 MZUSP Altamira Brazil -3.5039723 Rio Xingu 1 -52.03886 13 14 MZUSP 74687, 9 1 Para holotype 1412767350 [331,1327,1879,1904] MZUSP Santarem Brazil -2.4166667 Rio Tapajos 21 -54.733334 13 14 MZUSP 3528, 3 1 Para holotype 1412767336 MZUSP Oriximina Brazil -1.7666667 Rio Trombetas 21 -55.866665 13 14 MZUSP 5449, 4 1 Para holotype 1412767329 [665,1433,1952,1977] ANSP Terezina Brazil Rio Parnaiba 13 14 ANSP 84415, 5 1 Piaui holotype 1412767331 [151,1185,1988,2013] MZUSP Teresina Brazil -5.0833335 Rio Parnaiba 21 -42.833332 13 14 MZUSP 5091, 15 1 Piaui holotype 1412767338 MZUSP Teresina Brazil 14 15 Rio Parnaiba 13 14 MZUSP 74891, 14 1 Piaui holotype 1412767343 ANSP Russas Brazil Rio Jaguaribe 14 15 ANSP 84416, 2 1 Ceara holotype 1412767330 [245,934,188,213] Brazil Rio Araguaia 14 15 1 Goias holotype 1412767346 Brazil Britania -15.893222 Rio Vermelho 14 -52.25028 Rio Araguaia 14 15 1 Goias holotype 1412767348 MZUSP Aruana Brazil -15.173111 Rio Araguaia 1 -51.157528 14 15 MZUSP 4814, 28 1 Goias holotype 1412767333 [511,1035,260,285] MZUSP Brazil Rio Araguaia 14 15 MZUSP 54523, 1 1 Goias holotype 1412767345 MZUSP Minacu Brazil Tocantins 14 15 MZUSP 110039, 3 1 Tocantins holotype 1412767328 Brazil Balsas -7.5405555 Rio Balsas 21 -46.039165 rio Parnaiba 14 15 1 Maranhao holotype 1412767335 [431,1293,332,357] Paraguay Rio Mutuca 14 15 1 Mato Grosso holotype 1412767332 Paraguay -15.355556 Paraguai 21 -56.10111 14 15 1 holotype 1412767349 [274,1264,404,429] Paraguay -16.118334 Paraguai 21 -57.7425 14 15 1 holotype 1412767337 Paraguay -16.053778 Rio Cuiaba 1 -57.808834 Pocone 14 15 1 holotype 1412767341 Paraguay Canarana -17.847195 Rio Culuene 1 -57.43953 Rio Xingu 14 15 1 holotype 1412767326 Paraguay -13.517445 Rio Culuene 1 -53.078278 Canarana 14 15 1 holotype 1412767361 LIRP Barao de Melgaco Paraguay -13.494945 Rio Cuiaba 1 -53.082695 14 15 LIRP 713, 2 1 holotype 1412767356 MZUSP Paraguay -16.2 Rio Aripuana 21 -55.958336 Rio Madeira 14 15 MZUSP 101060, 1 1 holotype 1412767375 MZUSP Aripuana Paraguay -10.209167 Rio Aripuana 21 -59.460835 Rio Madeira 14 15 MZUSP 101069, 15 1 holotype 1412767364 Chapada dos Guimaraes Paraguay Rio Madeira Rio Cuiaba. Rio Aripuana. 14 15 1 holotype 1412767367 Paraguay Rio Cuiaba Rosario Oeste 14 15 1 holotype 1412767373 Paraguay Nova Brasilandia 14 15 1 holotype 1412767362 Paraguay Rio Manso. Mato Grosso do Sul 14 15 1 Rio Manso. Mato Grosso do Sul holotype 1412767371 [268,1246,764,789] Paraguay -18.428473 Paraguai 14 -54.83389 14 15 1 holotype 1412767351 Paraguay -56.24708 Aquidauana 1 -19.576029 14 15 1 holotype 1412767365 Paraguay -17.92111 Corumba 15 -57.475445 14 15 1 holotype 1412767366 Paraguay -17.51361 Pocone 15 -57.24064 14 15 1 holotype 1412767357 [470,1432,871,896] MZUSP Aquidauana Paraguay Rio Negro 14 15 MZUSP 59706, 21 1 Rio Negro holotype 1412767359 [151,871,908,933] Paraguay Parana 14 15 1 Parana holotype