Zootaxa 3994 (1): 133–141 www.mapress.com /zootaxa / Copyright © 2015 Magnolia Press Article ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3994.1.7 http://zoobank.org/urn:lsid:zoobank.org:pub:D9D78466-2E20-4BAA-8520-E0F2FBBDDB1D Bryconops munduruku (Characiformes: Characidae), a new species of fish from the lower Tapajós River basin, Brazil CÁRLISON SILVA-OLIVEIRA1, ANDRÉ LUIZ C. CANTO2 & FRANK RAYNNER V. RIBEIRO1,2 1 Programa de Pós-Graduação em Recursos Aquáticos Continentais Amazônicos, Instituto de Ciências e Tecnologia das Águas, Universidade Federal do Oeste do Pará. Rua Vera Paz, s/n – Sal, CEP 68035-110, Santarém, Pará, Brazil. E-mail: [email protected]; [email protected] 2 Coleção Ictiológica da Universidade Federal do Oeste do Pará. Campus Amazônia. Avenida Mendonça Furtado, 2.946 - Fátima, CEP 68040-470, Santarém, Pará, Brazil. E-mail: [email protected] Abstract A new species of Bryconops is described from a right tributary of the lower Tapajós River, State of Pará, Brazil. Bryconops munduruku, sp. nov., differs from its congeners, except B. inpai and B. piracolina, by having a black adipose fin (vs. adipose fin hyaline in alcohol) and, except B. inpai, by possessing two humeral blotches (vs. lack of humeral blotch or only one humeral blotch). Bryconops munduruku differs from B. inpai by having a uniform color pattern on the posterior portion of the side of the body (vs. a dark stripe extending posteriorly from the half of the anal-fin base onto the base of the middle caudal-fin rays). It differs further from B. inpai and B. piracolina by the presence of a black adipose fin that is hyaline along its base (vs. entirely black adipose fin in B. inpai and B. piracolina). The new species is allocated in the subgenus Creatochanes by having a maxillary bone with one to three teeth on both sides, and its posterior extension reaching the junction of second and third infraorbital bones. Key words: subgenus Creatochanes, taxonomy, Tapajós National Forest, Amazon basin Resumo Uma espécie nova de Bryconops descrita de um afluente da margem direita do baixo rio Tapajós, Pará, Brasil. Bryconops munduruku difere de seus congêneres, exceto B. inpai e B. piracolina, pela nadadeira adiposa preta (vs. nadadeira adiposa hialina em álcool) e, exceto B. inpai, por apresentar duas manchas umerais. Bryconops munduruku difere de B. inpai pelo padrão uniforme de coloração na porção posterior do corpo (vs. uma faixa lateral escura que se estende posteriormente da metade da base da nadadeira anal at a base dos raios medianos da nadadeira caudal). Difere ainda de B. inpai e B. piracolina pela nadadeira adiposa escura distalmente, mas hialina ao longo de sua base (vs. nadadeira adiposa completamente escura em B. piracolina e B. inpai). A espécie nova alocada no subgênero Creatochanes por apresentar de um a três dentes em cada osso maxilar e a margem posterior da maxila alcançando a margem posterior do segundo infraorbital. Introduction Bryconops was proposed by Kner (1858) to allocate two new species, Bryconops alburnoides Kner and Bryconops lucidus Kner (= Bryconops alburnoides Kner), described based on specimens from the Guaporé and Branco rivers, Brazil, respectively. Currently 19 valid species are recognized in the genus (Lima et al. 2003; Chernoff & Machado-Allison 2005; Wingert & Malabarba 2011; Eschmeyer 2015). The genus is restricted to freshwater drainages of South America, occurring in the Amazon, Orinoco, Tocantins, São Francisco, and Paraguay river basins, and in coastal rivers of northern Brazil and the Guianas (Lima et al. 2003; Lima & Caires 2011; Polaz et al. 2014; Ramos et al. 2014; Eschmeyer 2015). Chernoff & Machado-Allison (1999, 2005) divided the species of Bryconops into two subgenera, Bryconops and Creatochanes Günther 1864, based on the number of maxillary teeth and position of the posterior margin of the maxillary bone in relation to the contact between the second and third infraorbital bones. Accepted by M.R. de Carvalho: 18 May 2015; published: 30 Jul. 2015 133 Among the valid species, eight exhibit characters proposed for the subgenus Bryconops: B. alburnoides, B. caudomaculatus (Günther 1864), B. collettei Chernoff & Machado-Allison 2005, B. disruptus Machado-Allison & Chernoff 1997, B. durbini (Eigenmann 1908), B. gracilis (Eigenmann 1908), B. magoi Chernoff & MachadoAllison 2005, and B. piracolina Wingert & Malabarba 2011. Eleven species exhibit characters proposed for the subgenus Creatochanes: B. affinis (Günther 1864), B. colanegra Chernoff & Machado-Allison 1999, B. colaroja Chernoff & Machado-Allison 1999, B. cyrtogaster (Norman 1926), B. giacopinii (Fernández-Yépez 1950), B. humeralis Machado-Allison, Chernoff & Buckup 1996, B. imitator Chernoff & Machado-Allison 2002, B. inpai Knöppel, Junk & Géry 1968, B. melanurus (Bloch 1794), B. transitoria Steindachner 1915, and B. vibex MachadoAllison, Chernoff & Buckup 1996. In this paper, a new species of Bryconops is described from the igarapé Açu, a tributary of the lower Tapajós River basin in Brazil. Material and methods Methods for taking morphometric and meristic data followed Fink & Weitzman (1974) and Machado-Allison et al. (1993, 1996), except for counts of horizontal scale rows below the lateral line that were taken following Malabarba & Bertaco (1999). All measurements were taken point to point, on the left side of the body, with digital caliper to the nearest 0.1 mm, and are summarized in the species accounts and table as percentages of Standard Length (SL) or Head Length (HL). Drawings and counts of vertebrae, supraneurals, pterygiophores, procurrent caudal-fin rays, gill rakers on the first branchial arch, and branchiostegal rays were taken from cleared and stained (c&s) paratypes, prepared following the clearing and staining methods of Taylor & van Dyke (1985). Institutional abbreviations follow Ferraris Jr. (2007), with the addition of the Fish Collection of Universidade Federal do Oeste do Pará (UFOPA-I). Comparisons to species not listed in comparative material were based on the literature. In the description, counts are followed by number of frequencies, corresponding to the number of specimens analyzed; asterisk within parentheses (*) indicates the holotype count. In the list of comparative material examined, museum abbreviations and catalog numbers are followed by the total number of specimens in that lot, range of standard length, and collecting data. Bryconops munduruku, new species (Figures 1–3; Table 1) Holotype. INPA 46510, male, 76.6 mm SL, Brazil, Pará State, Aveiro, igarapé Açu, tributary of the right margin of the Tapajós River, about 10 km of the city of Aveiro, 0336’0.99’’S 5514’59.14’’W, 23 May 2013, C. Silva-Oliveira & S. R. Oliveira. Paratypes. All collected with holotype, Brazil, Pará State, Tapajós River basin: MCP 48315 (5, 34.0–79.6 mm SL); INPA 46511 (11, 33.5–91.7 mm SL; 2 c&s, 31.8–74.1 mm SL); UFOPA-I-00655 (7, 33.4–96.4 mm SL; 5 c&s, 47.6–63.3 mm SL). Diagnosis. Bryconops munduruku differs from its congeners, except B. inpai and B. piracolina, by possessing a black adipose fin (vs. adipose fin hyaline in alcohol). It differs from B. inpai and B. piracolina by possessing a hyaline band on the black adipose-fin base (vs. entirely black adipose fin in B. piracolina and B. inpai). Bryconops munduruku differs further from B. piracolina by having a hyaline dorsal fin (vs. presence of a large black blotch on the dorsal-fin base). Bryconops munduruku differs from other species of the subgenus Creatochanes, except B. inpai, by possessing two humeral blotches (vs. lack of humeral blotch or humeral region with a single humeral blotch in B. humeralis and B. vibex). It differs further from B. inpai due to the uniform color pattern on the posterior portion of the side of the body (vs. presence of a dark stripe extending posteriorly from the half of the anal-fin base onto the base of the middle caudal-fin rays). Description. Morphometric data presented in Table 1. Body shape compressed, greatest body depth located just anterior to dorsal-fin origin. Dorsal profile of body slightly convex from snout tip to dorsal-fin origin; straight to slightly concave from dorsal insertion to adipose-fin origin, gently sloping to beginning of caudal peduncle. 134 · Zootaxa 3994 (1) © 2015 Magnolia Press SILVA-OLIVEIRA ET AL. Caudal peduncle concave along dorsal and ventral margins. Ventral profile of head and body convex from lower lip to pelvic-fin origin, and straight from this point to anal-fin origin. Ventral profile straight and posterodorsally inclined along anal-fin base. TABLE 1. Morphometric data of holotype and 30 paratypes of Bryconops munduruku. Holotype is included in the ranges and means. SD = standard deviation. Standard Length (mm) Holotype Range Mean SD 76.6 31.8–96.4 61.4 - Percents of Standard Length Predorsal length 41.4 41.4–50.6 48.8 1.6 Preanal length 66.7 60.4–67.3 65.3 1.4 Prepelvic length 49.0 47.4–52.6 49.8 1.3 Prepectoral length 26.4 25.5–29.1 27.1 0.8 Dorsal-fin base length 13.1 9.8–17.9 14.5 1.4 Dorsal origin to pectoral origin 37.3 31.5–38.5 35.8 1.7 Dorsal origin to pelvic origin 33.7 27.1–35.3 31.9 2.0 Dorsal terminus to anal origin 35.1 27.6–37.8 34.1 2.5 Dorsal terminus to anal origin 28.1 21.5–30.8 27.4 2.2 Dorsal terminus to pelvic origin 33.4 27.7–35.4 32.1 1.9 Dorsal terminus to anal terminus 31.9 26.3–35.3 31.0 1.7 Dorsal terminus to adipose origin 22.1 21.3–26.6 22.9 1.1 Adipose terminus to hypural plate 14.6 11.7–16.0 14.4 1.1 Pectoral origin to pelvic origin 23.8 20.6–25.6 23.2 1.1 Anal-fin base length 25.7 24.6–29.5 27.2 1.2 Anal origin to adipose origin 30.4 27.1–33.0 30.1 1.6 Length of caudal peduncle 12.4 9.6–14.3 12.5 1.0 Adipose origin to base of last anal-fin ray 10.8 10.8–13.5 12.4 0.7 Maxillary length 14.0 13.1–15.7 14.1 0.7 Snout length 8.0 5.9–8.2 6.9 0.6 Horizontal orbit diameter 8.4 7.1–12.0 9.7 1.0 Head length 26.0 22.2–29.2 26.1 1.4 86.4 81.7–102.7 89.1 4.6 Percents of Head Length Tip of snout to tip supraoccipital spine Maxilla to pectoral origin 45.7 44.6–59.8 50.8 3.9 Maxilla to posterior margin of opercle 55.8 51.7–69.3 57.8 3.7 Anterior margin of orbit to maxilla 38.2 32.6–46.7 39.2 2.7 Posterior margin of orbit to opercle 41.0 31.3–48.7 36.7 3.9 Maxillary length 53.8 48.5–60.7 54.0 2.1 Snout length 30.7 21.5–32.0 26.3 2.2 Horizontal orbit diameter 32.2 32.2–44.7 7.5 2.8 Mouth terminal. Posterior extension of maxilla reaching junction between second and third infraorbital bones (Fig. 2); third infraorbital moderately developed, reaching preopercle ventrally. Supraorbital bone present. Premaxillary teeth in two rows, with midcentral cusp more developed than remaining cusps; outer teeth row with 5* (9) or 6 (3) tricuspid teeth; inner tooth row with 5* (10) pentacuspid teeth. Maxillary with 1 (2), 2* (15), or 3 (1) tri- or tetracuspid teeth on contralateral parts (Fig. 3). Dentary with 5 (2) or 6* (6) penta- or hexacuspid teeth, followed by 6 smaller conical teeth. NEW BRYCONOPS FROM THE TAPAJÓS RIVER Zootaxa 3994 (1) © 2015 Magnolia Press · 135 FIGURE 1. Bryconops munduruku, INPA 46510, holotype, male, 76.6 mm SL, Brazil, Pará State, Aveiro, Igarapé Açu, tributary of the lower Tapajós River basin. FIGURE 2. Bryconops munduruku, UFOPA-I-00655, paratype, 71.5 mm SL (c&s). Left side of the premaxilla, maxilla, and dentary. Lateral view. Scale bar = 1 mm. Dorsal-fin rays ii,8 (3) or ii,9* (28); first unbranched ray about one-half length of second unbranched ray. Dorsal-fin origin slightly ahead of mid-body, positioned vertically through anterior third of pelvic-fin base. Posterior margin of dorsal fin straight to slightly concave. Adipose-fin origin approximately at vertical through base of 18th to 19th anal fin branched rays. Principal caudal-fin rays i,8,8,i (7) or i,8,9,i* (23). Lobes of caudal fin unequal, lower lobe slightly longer than upper lobe. Dorsal procurrent caudal-fin rays 14 (5), ventral procurrent caudal-fin rays 13 (4) or 14 (1). Pectoral-fin rays i,10 (15) or i,11* (15). Tip of pectoral fin reaching slightly beyond half of distance between pectoral and pelvic fins. Pelvic-fin rays i,7*(30). Pelvic fin originating at vertical through middle of dorsal-fin base, reaching slightly beyond half the distance between pelvic and anal fins. Anal-fin 136 · Zootaxa 3994 (1) © 2015 Magnolia Press SILVA-OLIVEIRA ET AL. rays iii, 23* (16), iii, 24 (6), iii, 25 (6), or iii, 26 (2). Anal-fin origin located posterior to vertical through base of last dorsal-fin ray. Last unbranched anal-fin ray and first to fourth branched rays slightly longer than remaining anal-fin rays. Distal margin of anal fin concave. FIGURE 3. Bryconops munduruku,UFOPA-I-00655, paratype,63.3 mm SL(c&s). Maxilla (MA), infraorbitals (IO) 1-5 and antobital (AO). Scale bar = 1 mm. Lateral line reaching base of caudal-fin rays. Longitudinal scales in lateral line 44 (6), 45* (18), or 46 (8). All scales of lateral line pored. Scale rows between lateral line and dorsal-fin origin 7* (22) or 8 (8); scale rows between lateral line and pelvic-fin origin 3* (18) or 4 (12). Predorsal scales 10 (9) or 11* (5), arranged in a regular series. Scale rows around caudal peduncle 15 (5) or 16* (3). Precaudal vertebrae 19 (5); caudal vertebrae 22 (1) or 23 (4); total vertebrae 41 (1) or 42 (4). Supraneurals 7 (4). First dorsal fin pterygiophore located between 11th and 12th vertebrae. Gill rakers of first gill arch 16 (5): 2 (5) hypobranchial, 7 (5) ceratobranchial, 1 (5) on cartilage between ceratobranchial and epibranchial, 6 (5) epibranchial. Gill rakers setiform. Branchiostegal rays 5: 4 (5) on ceratohyal and 1 (2) on epihyal. Color in alcohol. Overall ground coloration of body dusky brown to yellowish. Dorsal profile of head and snout dark. Infraorbitals, maxillary, and ventral region of opercle silver. Infraorbital 5 and dorsal portion of opercle with scattered chromatophores. Lips and anterior portion of maxilla with dense concentration of chromatophores. Gular region light. Dorsolateral region of body darkened. Scales of lateral region of body with chromatophores concentrated on posterior borders. Two humeral spots present; first humeral spot conspicuous, vertically elongated, NEW BRYCONOPS FROM THE TAPAJÓS RIVER Zootaxa 3994 (1) © 2015 Magnolia Press · 137 extending over three rows of scales above lateral line and horizontally over two rows of scales; second humeral spot inconspicuous, vertically elongated; its form resembling the number 3. Dorsal-fin rays darkened from base to mid-section, light at distal portion. Black adipose fin, but hyaline along its base. Caudal fin with concentrated chromatophores on half nearest most external rays of ventral and dorsal lobes. Pectoral, pelvic, and anal fins hyaline, with few scattered chromatophores. Color in life. General body color reddish on dorsolateral region above lateral line; light grey from lateral line to ventral scale series. Upper portion of head, opercle, intraorbital bones, and maxilla dark grey. Upper margin of orbit red; anterior and posterior margins yellowish. Dorsal fin red from base to mid-length of rays; distally hyaline. Black adipose fin, with narrow hyaline band at base. Caudal fin red at base to mid-length of dorsal and ventral rays; distal region of lobes hyaline. Pectoral and pelvic fins, as well as first anal-fin rays, yellow. Sexual dimorphism. Mature males bear hooks on the anal, pelvic, and dorsal-fins rays. Dorsal and pelvic-fin hooks are smaller, fewer in number and spine-like. Anal-fin hooks larger and more numerous, present from distal half of the third unbranched ray, decreasing in number and size to15th branched ray. Hooks absent in females. Distribution. Bryconops munduruku is known from the igarapé Açu, a tributary on the right margin of the Tapajós River, about 10 km from Aveiro, Pará State, Brazil (Fig. 4). Etymology. The specific epithet is given in allusion to a tribe of Munduruku Indians denominated TapajósTapera, who settled on the right margin of the Tapajós River, giving rise to what today is the city of Aveiro, the type locality of Bryconops munduruku. A noun in apposition. Ecological notes. igarapé Açu presents clear water running over sandy beds. Its headwaters are located within the limits of the Tapajós National Forest and are densely covered by marginal vegetation. Representatives of Bryconops munduruku were collected in rapid flowing waters among stretches, where it occurs syntopically with Bryconops cf. giacopinii, Hyphessobrycon cf. agulha, Hemigrammus sp., and Moenkhausia comma. FIGURE 4. Map of part of Central and Northern South America, showing the type locality of Bryconops munduruku (black square) in the lower Tapajós River basin. 138 · Zootaxa 3994 (1) © 2015 Magnolia Press SILVA-OLIVEIRA ET AL. FIGURE 5. Igarapé Açu, type locality of Bryconops munduruku, lower Tapajós River, Aveiro, Pará State, Brazil. Discussion Bryconops munduruku shares with its congeners characters proposed by Chernoff & Machado-Allison (1999) as synapomorphies for the genus, as follows: (1) ventral margin of maxillary bone curved at an angle of approximately 90°; (2) infraorbital sensory canal well developed, reaching the antorbital bone; and (3) sensory canal extends onto nuchal scales. Within the genus Bryconops, Chernoff & Machado-Allison (1999) proposed the monophyly of each subgenera, Bryconops and Creatochanes, based on two putative synapomorphies. Thus, subgenus Bryconops is defined by the absence, or sporadic presence of a sole conical tooth located in a single maxillary bone, and the posterior end of the maxilla not reaching the junction of second and third infraorbital bones; and subgenus Creatochanes is recognized by the presence of one to three teeth on each maxilla, and the posterior end of the maxilla reaching the region of contact between the second and third infraorbital bones. As for Bryconops munduruku, it possesses the characters proposed by Chernoff & Machado-Allison (1999, 2005) that define the subgenus Creatochanes. Among its congeners, B. munduruku resembles B. inpai by having a deep body, presence of two humeral blotches, and a black adipose fin. However, B. munduruku can be readily distinguished from B. inpai by the uniform color pattern on the posterior portion of the body side (vs. a dark stripe on the posterior portion of the body side). Although Bryconops inpai was described from igarapé Barro Branco in the lower portion of the Negro River, this species also occur along the middle and upper Negro River in Brazil, and in the Casiquiare River in Venezuela (Lima et al. 2003). Analysis of Bryconops inpai holdings deposited at the Instituto Nacional de Pesquisas da Amazônia (INPA 31573) allowed us to extend the distributional range of this species to the Trombetas River, near Oriximin, Pará State, Brazil. NEW BRYCONOPS FROM THE TAPAJÓS RIVER Zootaxa 3994 (1) © 2015 Magnolia Press · 139 Bryconops munduruku is the fourth species of the genus described from the Tapajós River. Its congeners described from the Tapajós River are: B. durbini, B. gracilis, and B. transitoria. Even though the new species can be readily diagnosed among it congeners in the Tapajós River, the validity of B. transitoria and B. gracilis needs further investigation. Steindachner (1915) described B. transitoria and stressed its similarity with B. melanurus, but distinguished them based on morphometric accounts, and by the number of anal-fin rays (23–27 vs. 28–29, respectively). Bryconops gracilis was described based on a sole specimen, which was also recognized as similar to B. melanurus. We were not able to find any record of these two species in the Tapajós River. Comparative material examined Bryconops alburnoides: Brazil. INPA 12316 (1, 107.5 mm SL), Tocantins River; INPA 30701 (1, 145.7 mm SL), Jaú River; INPA 37794 (1, 81.1 mm SL), Negro River; INPA 36079 (6, 112.3–115.1 mm SL), Branco River. Bryconops caudomaculatus. Brazil. INPA 12397 (4, 69.3–77.1 mm SL), Trombetas River; INPA 12446 (2, 70.0– 73.0 mm SL), Jamari River. Bryconops durbini. Brazil. UFOPA-I-00337 (109, 18.2–61.0 mm SL), Lago Verde, Tapajós River. Bryconops aff. magoi: INPA 35801 (28, 27.1–38.0 mm SL), Purus River. Bryconops inpai: INPA 13249 (3, 63.2–72.9 mm SL), Urubu River; INPA 29524 (1, 66.7 mm SL), Trombetas River; INPA 31573 (1, 59.3 mm SL), Anebá River. Bryconops cf. vibex: INPA19638 (3, 49.8–66.3 mm SL), Negro River; INPA 19634 (6, 53.8–83.5 mm SL), Negro River; INPA 19636 (7, 31.3–79.6 mm SL), Negro River. Bryconops giacopinii: INPA 32636 (7, 57.1–67.7 mm SL), Negro River. Bryconops melanurus: Brazil. INPA 12443 (3, 91.1–91.9 mm SL), Caraipé River; INPA 26656 (1, 109.0 mm SL), Madeira River; INPA 38579 (3, 67.3–81.4 mm SL), Madeira River; UFOPA-I-00336 (16, 84.1–109.7 mm SL), Moju River; UFOPA-I-00338 (11, 49.1–78.0 mm SL), Curuá River. Acknowledgements The authors are indebted to the following: Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) for authorization to collect fishes (number 35649-2) in the Floresta Nacional do Tapajós; Programa de Pesquisas em Biodiversidade (PPBio/Amazônia Oriental), and Universidade Federal do Oeste do Pará (UFOPA) for financial and logistical support. We thank David de Santana for suggestions to the manuscript; Sérgio R. Oliveira for helping in the field and to Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the fellowship granted to Cárlison Silva-Oliveira. References Chernoff, B. & Machado-Allison, A. (1999) Bryconops colaroja and B. colanegra, two new species from the Cuyun and Caron drainages of South America (Teleostei: Characiformes). Ichthyological Exploration of Freshwaters, 10, 355–370. Chernoff, B. & Machado-Allison, A. 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