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Moenkhausia doceana (Steindachner, 1877) Figs. 1-5 Tetragonopterus doceanus Steindachner, 1877: 572-575. Type locality: rio Doce. Moenkhausia doceana (Steindachner): - Eigenmann, 1910: 437 (listed). - Eigenmann, 1917: 66; 73-75 (short redescription of species, but without analysis of syntypes). -Zarske & G��ry, 2001: 5-13 (comparison to M. margitae and data of some syntypes). -Zarske & G��ry, 2002: 52 (comparison to M. dorsinuda). - Lima et al., 2003: 148 (listed). -Bertaco & Lucinda, 2006: 65, 66 (comparison to M. pankilopteryx). - Lima et al., 2007: 58 (listed). Diagnosis. Moenkhausia doceana is promptly distinguished from all congeners [except M. affinis (Steindachner), M. barbouri Eigenmann, M. dorsinuda Zarske & G��ry, M. jamesi Eigenmann, M. justae Eigenmann, M. latissima Eigenmann, M. levidorsa Benine, M. margitae Zarske & G��ry, M. moisae G��ry, Planquette & Le Bail, M. newtoni, M. simulata (Eigenmann), M. tergimacula Lucena & Lucena] by a long anal fin, with 29-34 (mode 32) branched rays (vs. 28 or less in others species). Moenkhausia doceana differs from the aforementioned species, except M. tergimacula, by having 4-7 (mode 5) maxillary teeth (vs. 3 or less). Lastly, M. doceana differs of M. tergimacula by having 7-8 (mode 7) scale rows above lateral line and no marks anteriorly to dorsal-fin origin (vs. 5�� scales rows above lateral line and presence of a distinct dark, saddle-like mark on the body immediately anterior to dorsal-fin origin). Description. Morphometric data are summarized in Table 2. Body compressed, moderately long and high, greatest body depth at vertical through dorsal-fin origin. Dorsal profile of head convex from tip of upper jaw to vertical through anterior nostril; slightly concave or straight from that point to tip of supraoccipital spine. Dorsal profile of body slightly convex from posterior tip of supraoccipital spine to base of first dorsalfin ray, and straight that point to adipose-fin origin. Ventral profile of body convex from tip of lower jaw to pelvic-fin origin, straight or slightly convex from that point to anal-fin origin, and straight and posterodorsally slanted along of analfin base. Dorsal and ventral profile of caudal peduncle straight to slightly concave. Mouth terminal, premaxillary and dentary approximately the same size. Maxilla extending posteroventrally to first onethird of orbit, almost reaching tip of second infraorbital, approximately at 45 degrees angle relative to longitudinal axis of body. Main axis of maxilla straight, with approximately same width along entire length. Nostrils close to each other, anterior opening small and circular, posterior opening twice in size and reniform. Nostrils separated by skin flap when adpressed, almost covering the posterior nostril. Frontals not united anteriorly, with a triangle-shaped fontanel; parietal fontanel large, extending from epiphyseal bar to supraoccipital spine. Eyes relatively large, without adipose eyelid. Infraorbital bones complete, with six elements (Fig. 4). Laterosensory canal from first to sixth infraorbital close to inner margin of orbital rim. Third infraorbital largest, approximately twice the size of others (except second), with inferior margin slightly straight (Fig. 4), not contacting the laterosensory canal of preopercle ventrolaterally. Two tooth rows in the premaxilla: outer row with 4(13), 5(48), or 6(6) [4(4) or 5(2)] tricuspid teeth; inner row with 5(34) or 6(33) [5(6)], pentacuspid teeth. Dentary with 4(42) or 5(14) [4(3) or 5(3)] large, pentacuspid teeth, followed by a series of 10(1), 11(1), or 13(2) conical or triscupid teeth, abruptly smaller than the or five anterior largest teeth. Dorsal border of maxilla relatively straight. Maxilla with 4(12), 5(36), 6(15), or 7(7) [4(1), 5(3), or 6(2)] tri- or tetracuspid teeth along anteroventral margin (Fig. 5). Central median cusp of all teeth longer than remaining lateral cusps; cusps tips slightly curved inwardly on dentary and premaxillary teeth. Scales cycloid, with few radii (2-6), relatively small; circuli marked anteriorly and marginally (dorsal and ventral). Lateral line complete, extending from the superior portion of opercular opening to beginning of caudal fin with 34(2), 35(31), 36(29), 37(2), or 38(1) [34(2), 35(2), or 36(2)] perforated scales; horizontal scales rows between dorsal-fin origin and lateral line 7(23), 7��(10), or 8(37) [7(1) or 8(3)]; horizontal scales rows between lateral line and pelvic-fin origin 6(4), 6��(4), or 7(49) [6(1), 6��(1), or 7(2)]. Predorsal scales 10(20) or 11(13) [10(2) or 11(4)]. Scale sheath along anal-fin base 14(3), 15(2), 16(8), 17(4), 18(2), or 19(2) [15(1) or 16(1)] in a single row. Circumpeduncular scales 14(33) or 15(12) [14(4) or 15(1)]. Axillary scale present, covering two scales posteriorly, distinctly triangular-shaped, thinner on posterior tip. Dorsal fin rays ii,9(77) [ii,9(6)]; first unbranched ray approximately half the length of second one or shorter. Dorsalfin origin slightly forward on midbody, at vertical through anterior one-third of pelvic-fin origin. Base of last dorsal-fin ray at vertical through distal tip of longest pelvic-fin ray, anterior to origin of anal fin. Tip of longest ray of adpressed dorsal fin at vertical through base of first two branched analfin rays. First dorsal-fin pterygiophore inserted between neural spine of eighth and ninth precaudal vertebra. Adipose fin present, well-developed. Pectoral fin with i,11(2), 12(15), 13(51), or 14(3) [i,12(3) or i,14(3)]; longest ray extending to one-third of pelvic-fin origin. Pelvic fin with i,7(77) [i,7(6)] rays. Pelvic-fin origin located anterior to vertical through dorsal-fin origin and with small bony hooks in mature males, absent in females. Tip of longest pelvic-fin ray reaching up to second or third branched ray anal-fin ray when adpressed. Anal-fin rays iv,29(2), 30(1), 31(10), 32(26), 33(23), or 34(6) [31(3), 32(2), or 33(1)]. Caudal fin forked, lobes slightly rounded or pointed, upper lobe slightly smaller than lower lobe, with i,9/8,i(77) [i,9/8,i(6)] rays. Caudal fin scaled, scales over base and along lower caudal-fin lobe up to middle; upper lobe with few scales, extending to its anterior one-third; scales gradually decreasing posteriorly in size. Dorsal procurrent caudal-fin rays 9(1), 10(1), or 11(2), and ventral procurrent caudal-fin rays 7(2), or 8(2). Branchiostegal rays 4(4). First gill arch with 8(4) gill rakers on epibranchial, 1(4) between epibranchial and ceratobranchial, 10(3) or 11(1) on ceratobranchial, and 2(4) on hypobranchial. Denticles on gill rakers present. Precaudal vertebrae 15(4) and caudal vertebrae 19(2) or 20(2). Total vertebrae 34(2) or 35(2). Supraneurals 3(1) or 4(3), filiform, with dorsal portion expanded and some bifurcated dorsally. Color in alcohol. Overall ground coloration yellowish, with light brown thin stripe extending longitudinally along lateral line from vertical through dorsal-fin origin to caudal peduncle. Head brownish dorsally, paler towards sides of head. Dark chromatophores concentrated on distal margin of scales resulting in reticulated pattern. Mid-dorsal region darker than flanks. Humeral region with horizontally rounded dark spot immediately above lateral-line scales, extending about two scales vertically and 4-6 scales horizontally. A second faint blotch behind humeral spot, comma shaped. Area above hypural plate with faint dark spot, almost triangle shaped. First rays of pelvic and anal fins with dark chromatophores, resulting in darker fin border. Dorsal fin with scattered dark chromatophores, more concentrated on anterior one-half.Anal fin with scattered dark chromatophores.Adipose-fin contour with dark chromatophores. Remaining fins hyaline on tips, with scattered dark chromatophores between unbranched rays (Fig. 2). Color in life. The following description is based on the examination of freshly specimens recently caught along northern Esp��rito Santo and southern Bahia rivers. Ground color pale yellow to silver gray, darker dorsally. Sides of body somewhat silver colored anteriorly to light red posteriorly. Longitudinal inconspicuous brilliant stripe at level of lateral line, silvery colored in some specimens. Humeral region with a faint horizontally rounded spot, followed by a second comma shaped faint blotch. Eyes yellowish to whitish. Opercle and preopercle region silver colored; fins yellowish to translucent (Fig. 3). A faint brownish spot, almost triangle shaped on caudal peduncle. Sexual dimorphism. Bony hooks on anal and pelvic fins, and also anal-fin profile distinguish males from females in Moenkhausia doceana. Mature males with very small bony hooks on the segments of the last unbranched and all branched anal-fin rays (one pair of bony hooks on posterior surface of hemitrichia), absents in females. Males with anal-fin distal margin slightly straight (Fig. 2a) while females with anal-fin distal margin falcate anteriorly (Fig. 2b). Gill glands (sensu Burns & Weitzman, 1996) were not found macroscopically on first gill arch on both sexes. Distribution. Moenkhausia doceana occurs along coastal river systems, with northern limits corresponding to the rio Jo��o de Tiba, on extreme southern Bahia river basins. The species is recorded until southern limits at the rio Riacho, a small coastal basin south of rio Doce in Esp��rito Santo State, and to the West in the middle rio Doce lake systems, rio Doce basin (Fig. 6). Known populations of M. doceana inhabit areas of mild relief, as in lakes of the rio Doce valley and along coastal drainages between Esp��rito Santo and southern Bahia states. The western portion of its distribution is within Coastal Tablelands area, a wide region with elevations usually not passing 150 m, coinciding with the Cenozoic sediments of the Barreiras Group, well seen along the whole coastal distributional area. Despite its range distribution, no significant morphological and meristic differences were found among M. doceana populations, suggesting that the species corresponds to a single taxon in all coastal drainages where it occurs. Regarding global biogeography regionalization of freshwater systems, the pattern of distribution of M. doceana fits the Northeastern Mata Atl��ntica ecoregion (sensu Abell et al., 2008). Gerais (Ottoni et al., 2011), Reserva Biol��gica do C��rrego Grande, Esp��rito Santo (Sarmento-Soares & Martins-Pinheiro, 2013), and Reserva Biol��gica Sooretama, Esp��rito Santo (LMSS and RFMP, pers. obs.). Popular name. Moenkhausia doceana is known as lambari (Lima et al., 2003) or piaba, in northern Esp��rito Santo and southern Bahia states. Ecological notes. Moenkhausia doceana inhabits second and third order streams, about two to three meters wide and 0.5 to 1.6 meters deep, characterized by clear water and moderate to fast current, sandy bottom with gravel, and riparian surrounding vegetation. The fishes were captured in the water column, mainly near the surface. It is an invertivore that eats aquatic insect larvae (e.g., Chironomidae), nymphs (e.g., Ephemeroptera), and fragments of terrestrial arthropods (ants, beetles, and spiders). Moenkhausia doceana occurs syntopically with Characidium aff. fasciatum Reinhardt, Hoplias malabaricus (Bloch), Astyanax aff. intermedius Eigenmann, Astyanax aff. lacustris (L��tken), Hyphessobrycon bifasciatus Ellis, Trichomycterus pradensis Sarmento-Soares, Martins-Pinheiro, Aranda & Chamon, Corydoras nattereri Steindachner, Pimelodella aff. vittata (L��tken), Poecilia vivipara Bloch & Schneider, and Geophagus brasiliensis (Quoy & Gaimard) (Sarmento-Soares & Martins Pinheiro, 2013). Moenkhausia doceana was recorded in conservation units such as the Parque Estadual do Rio Doce (PERD), Minas Phylogenetic reconstruction. Phylogenetic analysis, according to implied weighting procedures (Mirande, 2009, 2010), with consensus of ���k��� = 10.07571802 (three trees) and ���k��� = 11.06353351 (three trees) values, resulted in one cladogram with 2263 steps, Consistency Index (CI) of 0.16 and Retention Index (RI) of 0.66. Moenkhausia doceana is included in the Tetragonopterinae of Mirande (2010). It is sister group of (Tetragonopterus argenteus (Gymnocorymbus ternetzi (Stichonodon insignis (Poptella paraguayensis, Stethaprion erythrurus)))). Synapomorphies for this clade are: ch. 201:1 - denticles on gill rakers absent [reversed in M. doceana]; ch. 287:1 - 18 or more branched anal-fin rays; ch. 314:0 - bony hooks on last pelvic-fin ray of adult males absent; ch. 341:1 - humeral spot horizontally-ovate. Autapomorphies for M. doceana are: ch. 22:0 - frontal fontanel present; ch. 63:0 - anterior region of third infraorbital not much expanded relative to posterior region of second infraorbital; ch. 66:0 - fourth infraorbital present and well developed; ch. 82:0 - dorsal end Syntypes Non-type specimensRange Mean Range Mean SD N Standard Length (mm) 57.8-83.8 71.1 30.1-81.4 54.8 - 73Percents of Standard LengthBody depth 41.0-45.7 44.0 37.0-45.8 41.9 2.0 73 Head length 24.6-27.3 26.1 16.6-30.9 26.7 1.9 73 Head depth 31.6-35.7 33.6 27.0-35.7 32.3 2.6 56 Predorsal length 51.3-52.6 52.0 48.9-54.4 51.0 1.3 73 Prepelvic length 42.1-45.0 44.1 40.9-48.5 44.6 1.5 73 Pelvic fin to anal distance 16.2-18.6 17.0 13.3-21.5 16.3 1.4 70 Caudal peduncle depth 10.3-11.6 10.7 7.8-11.8 9.8 0.8 73 Dorsal-fin base length 13.6-14.3 13.9 12.4-17.1 14.2 0.8 73 Anal-fin base length 41.0-44.0 42.2 37.4-44.2 41.0 1.5 73 Pectoral-fin length 20.5-24.0 22.4 21.0-25.5 23.0 0.9 72 Pelvic-fin length 17.7-20.3 19.2 16.3-22.1 19.2 1.0 72 Dorsal-fin length 29.5-34.2 31.9 28.4-35.2 32.2 1.7 71 Anal-fin length 20.0-23.3 21.7 18.4-25.3 22.2 2.0 61 Caudal peduncle length 9.9-11.7 10.8 9.3-13.4 11.0 0.9 73 Dorsal fin to adipose-fin distance 40.1-42.6 41.0 37.1-42.7 40.0 1.1 73 Eye to dorsal-fin origin 36.1-39.3 37.9 33.9-40.1 36.4 1.4 73 Dorsal origin to caudal origin 55.2-58.1 56.5 50.3-59.8 56.4 1.4 73Percents of head lengthInterorbital width 32.5-35.3 33.9 27.5-36.1 31.3 1.7 70 Snout length 20.8-24.9 23.1 21.9-29.3 24.8 1.9 65 Orbital diameter 39.9-44.0 42.3 40.6-48.5 44.6 1.9 55 Upper jaw length 45.7-48.5 47.7 41.3-49.8 47.3 1.9 41 of laterosensory canal of preopercle and suprapreopercle not overlapping anterodorsal process of opercle; ch. 109:1 - ventral margin of horizontal process of anguloarticular perpendicular to laterosensory canal of dentary from medial view; ch. 135:1 - two or more maxillary teeth; ch. 225:0 - abdominal ribs on anterior caudal vertebrae absent; ch. 281:0 - seven or fewer supraneurals; ch. 305:0 - anterior ventral procurrent caudal-fin rays paired, only distally fused; ch. 315:1 - bony hooks on first pelvic-fin ray of adult males present; ch. 340:1 - insertion of pterotic aponeurosis on a lobe situated dorsal to horizontal semicircular canal. Material examined. Syntypes. All from Brazil, rio Doce. Tetragonopterus doceanus, NMW 57389, 2(2), 57.8-70.0 mm SL. NMW 57591, 2(2), 83.3- 83.3 mm SL. MNHN 1913-135, 1(1), 68.4 mm SL. MHNH 1913-136, 1(1), 62.2 mm SL. Moenkhausia doceana, Non-type specimens (ordered by river basin, and munic��pio). All from Brazil. Esp��rito Santo State, rio Riacho basin. Munic��pio de Aracruz. MBML 2728, 4, 57.4-65.0 mm SL (ex-CIUFES 130078), limit between Aracruz and Linhares, Lagoa do Aguiar. Esp��rito Santo State, rio Doce basin. Munic��pio de Colatina. MBML 678, 2(1), 21.7-30.1 mm SL, mouth of rio Santa Joana. MBML 3391, 1, 34.3 mm SL, rio Santa Maria do Rio Doce. MBML 3590, 1(1), 61.0 mm SL, c��rrego Miracema on right side of road ES-080 from S��o Domingos do Norte to Colatina. MNRJ 37747, 1, 56.2 mm SL, same data as MBML 3590. Munic��pio de Itarana. MBML 763, 1, 26.4 mm SL, rio Santa Joana. Munic��pio de Linhares. MCP 42079, 1, 43.6 mm SL, rio Pequeno. MNRJ 9708, 1, not measured, Lagoa Juparan��. MZUSP 1461, 2, 74.6-80.0 mm SL, rio Doce. MZUSP 1560, 6, 52.6-63.4 mm SL, rio Doce. MZUSP 1622, 3, 47.7-75.7 mm SL, rio Doce. MZUSP 17065, 1, 53.0 mm SL, Lagoa do Aviso. MZUSP 17069, 1, 42.6 mm SL, Lagoa Nova. MZUSP 17073, 1, 75.4 mm SL, Lagoa Juparan��. MZUSP 75137, 11, 52.4-75.9 mm SL, rio Doce. ZUEC 3583, 4, 18.2-25.6 mm SL, c��rrego Rancho Alto, Reserva Florestal Vale do Rio Doce. Munic��pio de Pancas. MBML 1749, 2(1), 35.7-36.6 mm SL, c��rrego Ub��, tributary of rio Pancas. MBML 1836, 2(1), 49.6- 22.9 mm SL, c��rrego Floresta, tributary of rio Pancas on fazenda Floresta Lajinha. MBML 2330, 2(2), 40.0- 49.8 mm SL, c��rrego S��o Bento, tributary of rio Pancas in S��o Bento. Munic��pio de Santa Teresa. MZUSP 1562, 1, 64.2 mm SL, rio 25 de Julho. MZUSP 1629, 3, 57.6-62.8 mm SL, rio 25 de Julho. Minas Gerais State, rio Doce basin. Munic��pio de Frei Inoc��ncio. MCP 17771, 3, 30.5-30.9 mm SL, rio Sua��u�� tributary of rio Doce, under bridge on road BR-116. Munic��pio de Maril��ria. MZUSP 28972, 1, 58.9 mm SL. Lago Dom Helv��cio, rio Doce Valley. MZUSP 28981, 15, 55.7-66.7 mm SL, Lagoa Carioca, rio Doce Valley. MZUSP 36648, 2, 57.1-67.8 mm SL, Lagoa Carioca, rio Doce Valley. MZUSP 107342, 5, 61.1-72.7 mm SL, Lagoa Carioca, Parque Estadual Rio Doce. Esp��rito Santo State, rio Barra Seca basin. Munic��pio de Linhares. MBML 48, 5(3), 41.5- 60.4 mm SL, rio Ipiranga in Pontal do Ipiranga. MCP 17769, 4, 20.1-48.0 mm SL, Lagoa do Cupido on road BR-101, on northern limits of Reserva Biol��gica de Sooretama. MCP 17770, 1, 32.5 mm SL, coastal river on km 118 of road BR-101, near Reserva Biol��gica de Sooretama. Munic��pio de Sooretama. MBML 5114, 52(7), 59.2-81.4 mm SL, second pool on road ES-356 near bridge over rio Barra Seca, inside Reserva Biol��gica de Sooretama. MBML 5169, 5(2), 35.4-42.2 mm SL, rio Barra Seca under bridge on road ES-356 around the area of Reserva Biol��gica de Sooretama. MNRJ 5342, 7, 55.1-64.0 mm SL, rio Barra Seca, on road Linhares to S��o Mateus (in precary conservation state). MNRJ 17588, 22, 38.6- 64.6 mm SL, rio Barra Seca. MNRJ 40099, 10, 46.2-76.5 mm SL, c��rrego Cupido, near road BR-101 inside Reserva Biol��gica de Sooretama. MNRJ 40103, 1, 44.1 mm SL, c��rrego Parais��polis on its mouth in rio Barra Seca inside Reserva Biol��gica de Sooretama. MNRJ 40115, 6, 26.4-43.7 mm SL, rio Barra Seca under bridge on road ES-356 around the area of Reserva Biol��gica de Sooretama. MNRJ 40259, 54, 29.1-63.4 mm SL, same data as MBML 5169. Munic��pio de Jaguar��. MBML 5186, 1(1), 68.0 mm SL, rio Barra Seca on road between Nossa Senhora de F��tima and Vila Val��rio, around the area of Reserva Biol��gica de Sooretama. MNRJ 5324, 1, not measured, rio Cachimbau, on road Linhares to S��o Mateus. MNRJ 10916, 19, not measured, same data as MNRJ 5324. MN, Published as part of Carvalho, Fernando R., Sarmento-Soares, Luisa M. & Martins-Pinheiro, Ronaldo F., 2014, Redescription of Moenkhausia doceana (Steindachner, 1877) (Ostariophysi: Characiformes): a characid from the Northeastern Mata Atl��ntica ecoregion, Brazil, pp. 377-388 in Neotropical Ichthyology 12 (2) on pages 378-383, DOI: 10.1590/1982-0224-20140006, http://zenodo.org/record/4639092, {"references":["Steindachner, F. 1877. Die Susswasserfische des sudostlichen Brasilien (III). Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Classe, 74: 559 - 694.","Eigenmann, C. H. 1910. Catalogue of the fresh-water fishes of tropical and south temperate America. In: Scott, W. B. (Ed.). Reports of the Princeton University Expeditions to Patagonia 1896 - 1899. Princeton, N. J., 3: 375 - 511.","Eigenmann, C. H. 1917. The American Characidae - I. 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Phylogeny of the family Characidae (Teleostei: Characiformes): from characters to taxonomy. Neotropical Ichthyology, 8: 385 - 568.","Abell, R., M. L. Thieme, C. Revenga, M. Bryer, M. Kottelat, N. Bogutskaya, B. Coad, N. Mandrak, S. L. Balderas, W. Bussing, M. L. J. Stiassny, P. Skelton, G. R. Allen, P. Unmack, A. Naseka, R. Ng, N. Sindorf, J. Robertson, E. Armijo, J. Y. Higgins, T. J. Heibel, E. Wikramanayake, D. Olson, H. L. Lopez, R. E. Reis, J. G. Lundberg, M. H. S. Perez & P. Petry. 2008. Freshwater ecoregions of the world: a new map of biogeographic units for freshwater biodiversity conservation. Bioscience, 58: 403 - 414.","Ottoni, F. P., A. Q. Lezama, M. L. Triques, E. N. Fragoso-Moura, C. C. T. Lucas & F. A. R. Barbosa. 2011. Australoheros perdi, new species (Teleostei: Labroidei: Cichlidae) from the lacustrine region of the Doce River Valley, southeastern Brazil, with biological information. Vertebrate Zoology, 61: 137 - 145.","Sarmento-Soares, L. M. & R. F. Martins Pinheiro. 2013. A fauna de peixes na REBIO Corrego Grande e seu entorno direto, Espirito Santo, Brasil. Boletim do Museu de Biologia Mello Leitao, 31: 25 - 57.","Mirande, J. M. 2009. Weighted parsimony phylogeny of the family Characidae (Teleostei: Characiformes). Cladistics, 2009: 573 - 613."]}

Moenkhausia doceana: MCP 17990 (15, 39.3-49.1 mm SL), rio Bamburral, S��o Mateus, Esp��rito Santo, Brazil., Published as part of Vinicius A. Bertaco & Paulo H. F. Lucinda, 2006, Moenkhausia pankilopteryx, new species from rio Tocantins drainage, Brazil (Ostariophysi: Characiformes, Characidae)., pp. 57-68 in Zootaxa 1120 on page 66