Integrative taxonomy reveals two new cryptic species of Hyphessobrycon Durbin, 1908 (Teleostei: Characidae) from the Maracaçumé and middle Tocantins River basins, Eastern Amazon region Author Guimarães, Erick Cristofore 67E9AADF-0B92-437E-BB31-6352FBC3F42D Universidade Federal do Maranhão, Programa de Pós-Graduação em Biodiversidade e Biotecnologia da Amazônia Legal. Av. dos Portugueses 1966, Cidade Universitária do Bacanga, CEP 65080 - 805, São Luís, MA, Brazil. & Universidade Federal do Maranhão, Departamento de Biologia, Laboratório de Genética e Biologia Molecular, Av. dos Portugueses 1966, Cidade Universitária do Bacanga, CEP 65080 - 805, São Luís, MA, Brazil. erick.ictio@yahoo.com.br Author Brito, Pâmella Silva de EBBAE3D2-5BB2-4D2A-8A0D-B2B7CC29DC4F Universidade Federal do Maranhão, Programa de Pós-Graduação em Biodiversidade e Biotecnologia da Amazônia Legal. Av. dos Portugueses 1966, Cidade Universitária do Bacanga, CEP 65080 - 805, São Luís, MA, Brazil. & Universidade Federal do Maranhão, Departamento de Biologia, Laboratório de Genética e Biologia Molecular, Av. dos Portugueses 1966, Cidade Universitária do Bacanga, CEP 65080 - 805, São Luís, MA, Brazil. pamellabrito@hotmail.com Author Bragança, Pedro Henrique Negreiros 59164A3E-8380-4672-A39C-C2FE563F3C4F South African Institute for Aquatic Biodiversity, Private Bag 1015, Grahamstown, 6140, South Africa. pedrobra88@gmail.com Author Santos, Jadson Pinheiro 7248FF02-D740-4E2D-A7E0-6A9E81F76974 Universidade Federal do Maranhão, Programa de Pós-Graduação em Biodiversidade e Biotecnologia da Amazônia Legal. Av. dos Portugueses 1966, Cidade Universitária do Bacanga, CEP 65080 - 805, São Luís, MA, Brazil. & Universidade Estadual do Maranhão, Laboratório de Ictiofauna e Piscicultura Integrada, Centro de Ciências Agrárias, Campus Paulo VI, avenida Lourenço Vieira da Silva, n. 1000, bairro Jardim São Cristóvão, CEP 65055 - 310, São Luís, MA, Brazil. jadsonpesca@gmail.com Author Katz, Axel Makay 31E0031F-15ED-45B3-A0BB-CF985043D3B5 Universidade Federal do Rio de Janeiro, Laboratório de Sistemática e Evolução de Peixes Teleósteos, Departamento de Zoologia, Instituto de Biologia, Cidade Universitária, CEP 21941 - 599. Rio de Janeiro, RJ, Brazil. axelmk@gmail.com Author Costa, Luis Fernando Carvalho 2AE1615C-3F76-4867-9D7E-2942043F3D06 Universidade Federal do Maranhão, Departamento de Biologia, Laboratório de Genética e Biologia Molecular, Av. dos Portugueses 1966, Cidade Universitária do Bacanga, CEP 65080 - 805, São Luís, MA, Brazil. lfecc@yahoo.com.br Author Ottoni, Felipe Polivanov 6AB34592-DA6D-4AB7-88D8-1F63A97A12E3 Universidade Federal do Maranhão, Programa de Pós-Graduação em Biodiversidade e Biotecnologia da Amazônia Legal. Av. dos Portugueses 1966, Cidade Universitária do Bacanga, CEP 65080 - 805, São Luís, MA, Brazil. fpottoni@gmail.com text European Journal of Taxonomy 2020 2020-11-09 723 77 107 journal article 10.5852/ejt.2020.723.1145 2118-9773 4271695 10B6A707-DDA2-4D2F-9087-394933776459 Hyphessobrycon geryi Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. urn:lsid:zoobank.org:act: 75667753-6AB4-4E18-BBD6-3C33ECE68CD4 Figs 4–6 , Table 3 Morphological diagnosis (PAA) Hyphessobrycon geryi Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. clearly differs from most of its congeners, except members of the “Rosy tetra” clade, by the presence of a dark brown or black blotch on the dorsal fin (vs absence) and the absence of a midlateral stripe on the body (vs presence). The new species differs from most of its congeners from the “Rosy tetra” clade by possessing a conspicuous small slightly vertical elliptical humeral spot ( Fig. 1 ) (vs inconspicuous vertically elongated) humeral spot in Hyphessobrycon bentosi Durbin, 1908 , H. caru Guimar ães, De Brito, Feitosa, Carvalho-Costa, Ottoni, 2018 , H. hasemani Fowler, 1913 , H. piorskii Guimar ães, De Brito, Feitosa, Carvalho-Costa, Ottoni, 2018 ; approximately rounded humeral spot in H. erythrostigma (Fowler, 1943) , H. jackrobertsi Zarske, 2014 , H. minor Durbin, 1909 , H. pando Hein, 2009 , H. paepkei Zarske, 2014 , H. pyrrhonotus Burgess, 1993 , H. roseus (Géry, 1960) , H. socolofi Weitzman, 1977 , and H. sweglesi (Géry, 1961) ; humeral spot horizontally or posteriorly elongated in H. epicharis Weitzman & Palmer, 1997 , H. khardinae Zarske, 2008 , and H. werneri Géry & Uj, 1987 ; large vertical conspicuous humeral spot at least on males in H. eques (Steindachner, 1882) , H. haraldschultzi Travassos, 1960 , H. micropterus (Eigenmann, 1915) , H. megalopterus (Eigenmann, 1915) , H. simulatus (Géry, 1960) and H. takasei Géry, 1964 ; and absence of humeral spot in H. compressus (Meek, 1904) , H. dorsalis Zarske, 2014 , H. georgettae Géry, 1961 , H. pulchripinnis Ahl, 1937 , and H. rosaceus Durbin, 1909 ( Guimar ães et al. 2019 : fig. 2). The new species herein described differs from H. copelandi Durbin, 1908 , H. haraldschultzi Travassos, 1960 by the number of lateral series of scales (34–37 vs 29–31, 28–30 lateral line scales, respectively); from H. frickei Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. by the number of horizontal scale rows between lateral line and pelvic-fin origin (4–5, modally 4 vs 3); presence of conspicuous dark chromatophores on opercular zone (vs absence); presence of conspicuous dark chromatophores on the region posterior to humeral spot, extending to the end of caudal peduncle (vs absence); presence of a thin vertical line, formed by the concentration of dark chromatophores, at middle of humeral spot, extending one to two scales above and below the spot (vs absence); and dorsal-fin base with conspicuous pigmentation (vs less pigmented) (see Figs 1 , 4–5 ). Molecular diagnosis (CBB) Hyphessobrycon geryi Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. is a member of the H. copelandi clade, possessing the following 25 nucleotide substitutions: COI 90 (C+A), COI 126 (A+G), COI 138 (C+T), COI 189 (C+T), COI 192 (T+A), COI 237 (C+T), COI 264 (T+C), COI 282 (C+T), COI 285 (C+A), COI 312 (T+C), COI 384 (C+T), COI 402 (A+G), COI 429 (A+G), COI 435 (A+G), COI 486 (T+C), COI 522 (A+G), COI 525 (C+T), COI 547 (C+T), COI 582 (T+C), COI 621 (T+C), COI 624 (A+G), COI 678 (T+C), COI 684 (A+C), COI 690 (T+C), COI 696 (A+G). In addition, it differs from the other species of this group by the following eight nucleotide substitutions: COI 225 (T+C), COI 228 (A+C), COI 321 (G+A), COI 522 (G+A), COI 534 (G+A), COI 543 (A+G), COI 589 (C+T), COI 615 (G+A). Fig. 3. Geographical distribution of Hyphessobrycon frickei Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. and H. geryi Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. Red star denotes holotype and blue circle denotes paratypes of H. frickei Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. whereas white star denotes holotype and the black circles denote paratypes of H. geryi Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. Etymology The new species is named after the ichthyologist Jacques Géry (in memoriam) in recognition of his scientific contribution on Characiformes . Fig. 4. Hyphessobrycon geryi Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. , holotype (CICCAA 02403), 28.3 mm SL; Brazil: Maranh ão State: Lower Tocantins River basin. Fig. 5. Hyphessobrycon geryi Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. , paratype (CICCAA 02348), 22.7 mm SL; Brazil: Maranh ão State: Lower Tocantins River basin; living specimen photographed immediately after collection. Material examined Holotype BRAZIL • 27.8 mm SL; Pará State , Marabá municipality, Flexeiras River , Tocantins River basin ; 5°15′38″ S , 48°59′35″ W ; 20 Sep. 2016 ; E.C. Guimarães and C.H. Costa ; CICCAA 02403 Paratypes BRAZIL – Pará State • 1 spec. ; 25.6 mm SL; same collection data as for holotype; CICCAA 00722 • 13 specs ; 15.9–19.5 mm SL; Marabá municipality, Flexeiras River , Tocantins River basin; 5°15′38″ S , 48°59′35″ W ; 21 Sep. 2018 ; E.C. Guimarães and P.S. Brito leg.; CICCAA 02358 • 11 specs ; 16.9–19.2 mm SL; Marabá municipality, Flexeiras River , Tocantins River basin; 5°15′38″ S , 48°59′35″ W ; 9 Mar. 2018 ; E.C. Guimar ães and P.S. Brito leg.; CICCAA 02359 • 11 specs ; 18.8–22.2 mm SL; Bom Jesus do Tocantis , Jacundá River , Tocantins River basin ; 5°15′1″S , 48°48′58″ W ; 21 Dec. 2017 ; E.C. Guimarães and P.S. Brito leg.; CICCAA 02167 • 1 spec. ; 23.6 mm SL; Bom Jesus do Tocantis , Jacundá River , Tocantins River basin; 5°15′1″ S , 48°48′58″ W ; 21 Aug. 2017 ; E.C. Guimarães and P.S. Brito leg.; CICCAA 2348 • 5 specs ; 21.1–23.1 mm SL (C&S); Marabá municipality, Flexeiras River , Tocantins River basin; 5°15′38″ S , 48°59′35″ W ; 21 Aug. 2017 ; E.C. Guimarães and P.S. Brito leg.; CICCAA 04800 • 1 spec. ; 17.9 mm SL; Bom Jesus do Tocantins , Jacundá River , Tocantins River basin; 5°15′1″ S , 48°48′58″ W ; 21 Sep. 2018 ; E.C. Guimarães and P.S. Brito leg.; UFRJ 6919 • 4 specs ; 16.3–19.2 mm SL; Marabá municipality, Tocantins River basin; 5°15′38″ S , 48°59′35″ W ; 10 Jul. 2017 ; E.C. Guimarães and P.S. Brito leg.; CICCAA 01839 • 5 specs ; 18.0– 18.9 mm SL (C&S); Marabá municipality, Tocantins River basin; 5°15′38″ S , 48°59′35″ W ; 21 Sep. 2018 ; E.C. Guimarães and P.S. Brito leg.; CICCAA 02402 – Maranhão State • 1 spec. ; 26.1 mm SL; Igarapé Papagaio , Tocantins River basin; 5°8′59″ S , 48°22′33″ W ; 20 Jan. 2014 ; E.C. Guimarães leg.; CICCAA 01958 . Type locality Brazil , Pará State , Marabá municipality, Flexeiras River, Tocantins River basin, 5°15′38″ S , 48°59′35″ W ( Fig. 3 ). Description Morphometric data of holotype and paratypes are presented in Table 3. BODY. Small (with maximum SL of 23.6 mm ), compressed, moderately deep, greatest body depth slightly anterior to dorsal-fin base; body profile straight and downward directed from end of dorsal fin to adipose fin, straight or slightly convex between latter point and origin of dorsal most procurrent caudal-fin ray; dorsal profile of head convex from upper lip to vertical through eye; predorsal profile of body roughly straight, dorsal-fin base slightly convex, posteroventrally inclined; ventral profile of head convex from lower jaw to pelvic-fin origin; straight and posterodorsally slanted along anal-fin base; and slightly concave on caudal peduncle; jaws equal, mouth terminal. Maxilla reaching vertical to anterior margin of pupil. TEETH. Premaxillary teeth in two rows. Outer row with two tricuspid teeth; inner row with 5(2), 6(3) tricuspid teeth. Maxilla with 2(2), 3(2) or 4(1) pentacuspid teeth. Dentary with five (3) or six (2) larger tricuspid teeth followed by five (4) or six (1) smaller pentacuspid tooth ( Fig. 6 ). SCALES. Scales cycloid, three to eight radii strongly marked, circuli well-marked anteriorly, weaklymarked posteriorly; lateral line incompletely pored, with 7(31)* or 8(21) perforated scales. Longitudinal scales series including lateral-line scales 34(4), 35(2), 36(43)* or 37(1). Longitudinal scales rows between dorsal-fin origin and lateral line 5(46) or 6(4). Horizontal scale rows between lateral line and pelvic-fin origin 4(50) or 5(2). Scales in median series between tip of supraoccipital spine and dorsal-fin origin 9(35) or 10(15). Circumpeduncular scales 12(52). FINS. Dorsal-fin origin at midbody. Base of last dorsal-fin ray at vertical through first third of anal fin. Dorsal-fin rays ii + 9(50)* or iii + 9(2). First dorsal-fin pterygiophore main body located behind neural spine of 4 th vertebrae. Adipose fin present. Anal-fin origin aligned with vertical line through middle of dorsal fin, between 6 th and 8 th dorsal-fin rays. Anteriormost anal-fin pterygiophore inserting posterior to haemal spine of 11 th vertebrae. Anal-fin origin aligned with vertical line through middle of dorsal fin (between base of 6 th and 8 th dorsal-fin rays). Anal fin iii + 22(40)* or iii + 23(12); Anterior anal-fin margin slightly convex, with anteriormost rays more elongate and slightly more thickened than remaining rays, forming a distinct lobe. Remaining rays smaller with straight distal margin. Pectoral-fin rays 12(52) total rays. Tip of pectoral fin surpassing pelvic-fin base. Pelvic-fin rays 8(52) total rays, surpassing anal-fin origin. Caudal fin forked, upper and lower lobes similar in size. Principal caudal-fin rays 11+10(30) or 10+9(22); dorsal procurrent rays 8(8) or 9(2) and ventral procurrent rays 7(3) or 8(7). OSTEOLOGICAL COUNTS. Branchiostegal rays 10(10). First gill arch with 1(8), 2(2) hypobranchial, 11(2), 12(8) ceratobranchial, 1(10) cartilage between ceratobranchial and epibranchial, and 5(2) or 6(8) epibranchial gill-rakers. Supraneurals 4(8) or 5(2). Precaudal vertebrae 11 (10) and caudal vertebrae 19 (10). Total vertebrae 30(10). Coloration in alcohol ( Fig. 4 ) Ground coloration light yellowish brown. Humeral spot conspicuous, slightly vertically elliptical, with a thin vertical line, formed by the concentration of dark chromatophores, at middle of humeral spot, extending one to two scales above and below humeral spot. Flank with conspicuous chromatophores scattered mainly on middle region, from just after humeral spot, reaching caudal peduncle end. Ventral region lacking dark brown chromatophores. Dark brown chromatophores present on head, more concentrated on dorsal portion, tip of snout, and opercular region, becoming sparser on cheek. Dorsal-fin ground coloration hyaline, with conspicuous black or dark brown spot located on anterior portion of fin, reaching about 6 th ray, approximately between half to two thirds of fin depth. Conspicuous pigmentation on dorsal-fin base formed by concentration of conspicuous chromatophores. Anal and caudal fins hyaline. Anal and caudal fin with darker, usually dark brown, posterior margin. Adipose fin hyaline to light brown, with dark chromatophores at margin and posterior portion. Pectoral and pelvic fins hyaline; pelvic fin with variable amounts of dark brown pigmentation remaining depending on specimen preservation state. Fig. 6. Hyphessobrycon geryi Guimar ães, Brito, Bragança, Katz & Ottoni sp. nov. (CICCAA 02402), 23.2 mm SL. Jaw suspensory. A . Premaxillary. B . Maxilla. C . Dentary. Scale bar: 1 mm Color in life ( Fig. 5 ) Color pattern similar to coloration of preserved specimens, with more translucent body. Ground coloration translucent, white to grey, with orange pigmentation on vertebrae region, and usually with orange chromatophores. Ventral region anterior to anal-fin origin lighter. Chromatophores on humeral region black. Head darker than flank, especially on opercular and dorsal regions; ventrally lighter. Conspicuous black spot on dorsal fin, with yellow or white pigmentation on dorsal and ventral margins of spot; rest of dorsal fin hyaline.Anal fin base with light red chromatophores, with different degrees of intensity, with milk white pigmentation on anterior tip of anal fin, reaching between first to second rays. Adipose fin hyaline, with light red chromatophores mainly at base, and black chromatophores at margin. Pectoral and pelvic fins hyaline, with some sparser dark brown chromatophores, more concentrated at pelvic fin, and with milk white pigmentation on anterior tip of pelvic fin. Caudal fin with red pigmentation on almost fin, with inconspicuous light brown margin. Sexual dimorphism Bony hooks on fins is absent in all examined specimens. According to Malabarba & Weitzman (2003) the presence of bony hooks is a common dimorphic feature among characids species. Although this sexual dimorphism is not observed in all characid species, as in case of species described here. Color pattern is not sexually dimorphic either.