Hoplias lacerdae Miranda-Ribeiro, 1908

Oyakawa, Osvaldo T. & Mattox, George M. T., 2009, Revision of the Neotropical trahiras of the Hoplias lacerdae species-group (Ostariophysi: Characiformes: Erythrinidae) with descriptions of two new species, Neotropical Ichthyology 7 (2), pp. 117-140 : 134-137

Hoplias lacerdae Miranda-Ribeiro, 1908 Fig. 10

Hoplias lacerdae Miranda-Ribeiro, 1908: 3 [original description, type locality: rio Ribeira de Iguape, Iporanga, São Paulo state, Brazil; Holotype: MNRJ 211, dried specimen]. - Eigenmann, 1909: 374 [reference, occurrence in the rio Ribeira de Iguape]. - Eigenmann, 1910: 448 [reference, occurrence in the rio Ribeira de Iguape]. - Fowler, 1950: 364 [in species list]. - Azevedo et al., 1965: 103 [redescription, differences from H. malabaricus ]. - Britski, 1972: 81-82 [in part, species list, differences from H. malabaricus , occurrence in the rio Ribeira de Iguape basin]. - Géry et al., 1987: 368 [identification key]. - Bizerril, 1994: 56 [in part, occurrence in eastern river basins]. -Bizerril & Lima, 2000: 107 [occurrence in the rio Ribeira de Iguape]. - Oyakawa, 2003: 239 [species list, distribution]. - Oyakawa et al., 2006: 166 [species list with pictures, differences from H. malabaricus ]. -Oyakawa & Netto-Ferreira, 2007: 64 [species list, distribution]. - Menezes et al., 2007: 153 [typelocality, distribution].

Hoplias aff. lacerdae : - Malabarba, 1989: 127 [remarks on occurrence in the rio Jacuí, Rio Grande do Sul]. - Zaniboni Filho et al., 2004: 104 [biological and ecological features, with picture].

Hoplias lacertae : - Géry, 1977: 102 [unavailable name, lapsus calami, remarks on validity of name, identification key].

Holotype. BRAZIL. São Paulo State: MNRJ 211 , 750.0 mm SL, rio Ribeira de Iguape, Iporanga , no date, R. Krone. ( Fig. 11 ).

Material examined. BRAZIL. São Paulo State. Ribeira de Iguape basin: MNRJ 11217, 1, 443.0 mm SL, rio Juquiá, tributary to rio Ribeira de Iguape, Registro; MZUSP 7900, 1, 625.0 mm SL; MZUSP 7901, 1, 380.3 mm SL, rio Ribeira de Iguape; MZUSP

38569, 1, 157.2 mm SL, rio Ribeira de Iguape, Registro; MZUSP 7902, 1, 367.0 mm SL; MZUSP 37989, 2, 279.4- 417.2 mm SL, rio Pardo on the road Iporanga-Barra do Turvo, 14 km from Barra do Turvo; MZUSP 45433, 1, 234.9 mm SL, rio Ribeira de Iguape by rio Pilões, Eldorado; MZUSP 51104, 1, head length 244.5 mm SL, rio Juquiá, Registro; MZUSP 61748, 1, 150.5 mm SL, rio André Lopes, Iporanga; MZUSP 72953, 1, 388.6 mm SL, rio Jacupiranga, Registro. Santa Catarina State. Uruguay basin: MCP 12339, 1, 202.0 mm SL; MCP 12365, 1, 65.3 mm SL, rio Canoas between Vargem and São José do Cerrito, Campos Novos; MCP 12731, 1, 295.5 mm SL; MCP 12732, 1, 224.53 mm SL; MCP 12879, 2, 130.7-245.0 mm SL, rio Canoas at Passo do Canoas, SC-458, Campos Novos; MCP 12338, 1, 218.8 mm SL; MCP 12745, 2, 242- 249.3 mm SL; MCP 13143, 1, 152.9 mm SL; MCP 12878, 3, 297.2- 344.4 mm SL, rio Canoas, on Abdon Batista-Anita Garibaldi road, Campos Novos; MCP 12884, 1, 387.8 mm SL; MCP 12898, 3, 63.4-147.1 mm SL; MCP 41003, 1, 115.3 mm SL, rio Canoas, on Abdon Batista-Anita Garibaldi road; MCP 13420, 1, 180.0 mm SL; MCP 18427, 1, 203.8 mm SL; MCP 18828, 1, 226.6 mm SL; MCP 18925, 2, 242.6-301.0 mm SL; MCP 19151, 1, 402.1 mm SL; MCP 19035, 2, 243.1- 325.6 mm SL, rio do Peixe in Volta Grande, Concórdia; MCP 19146, 1, 411.5 mm SL; MCP 41004, 1, 253.3 mm SL, rio Riacho Grande, on Piritiba-BR 153 road, Concórdia; MCP 19147, 1, 415.1 mm SL, rio Uruguai in Itá; UFRGS 7530, 1, 238.6 mm SL, rio Jacutinga near water capture, Concórdia. Rio Grande do Sul State. Uruguay basin: INPA 26892, 1, 144.1 mm SL; MCP 12719, 3, 130.5- 185.3 mm SL, arroio Passo Alto, São Nicolau; MCP 10896, 3, 110.0- 207.7 mm SL, rio Garupá, tributary to rio Quaraí, on Uruguaiana-Quaraí road; MCP 11226, 1, 142.3 mm SL, tributary to rio Quaraí-Mirim on Quaraí-Alegrete road, Alegrete; MCP 11231, 4, 120.0- 179.9 mm SL; MNRJ 30393, 1, 145.7 mm SL, rio Garupá on Alegrete-Quaraí border; MCP 12089, 1, 288.8 mm SL; MCP 13421, 1, 301.1 mm SL; MNRJ 30394, 1, 302.1 mm SL, rio Ligeiro, on Marcelino Ramos- Maximiliano de Almeida road; MZUSP 37254, 1, 107.6 mm SL; MZUSP 43683, 1, 196.3 mm SL, rio Garupá, tributary to rio Quaraí; MCP 12674, 1, 284.3 mm SL, arroio Canoin on Pirapó- São Nicolau road, São Nicolau, Rio Grande do Sul; MCP 13210, 1, 285.8 mm SL, mouth of rio Ijuí-Mirim, Pirapó; MCP 14228, 1, 28.4 mm SL, tributary to rio Negro on Bagé-Aceguá road; MCP 14280, 1, 332.2 mm SL, arroio Ibicuí da Faxina, Santana do Livramento; MCP 16802, 2, 304.6- 349.2 mm SL, rio Ijuizinho near Cermissões dam; MCP 18875, 1, 250.3 mm SL, rio Uruguai in Marcelino Ramos; MCP 16818, 276.8 mm SL, rio Buricá near CERTHIL dam, Independência; MCP 18924, 1, 251.4 mm SL, rio Uruguai in Marcelino Ramos; MCP 19385, 2, 86.0- 124.6 mm SL, rio Canoas, on Tupitinga-Celso Ramos road, Tupitinga; MCP 19378, 2, 288.4- 309.4 mm SL, rio Pelotas at Passo da Pedra Oveira, Esmeralda; MCP 21650, 1, 170.8 mm SL, rio Quaraí-Mirim, on Uruguaiana-Quaraí road; MCP 26972, 2, 43.0- 43.7 mm SL, rio Ibirapuitã, under bridge upstream from the Biological Reserve of Ibirapuitã, Alegrete; MCP 27452, 1, 360.5 mm SL, Arroio Lagoão, rio Ibicuí drainage, on Santiago-Manoel de Freitas road; MCP 27481, 1, 144.4 mm SL, rio Itu, tributary to rio Ibicuí, Santiago; UFRGS 8394, 6, 68.8-94.9 mm SL, arroio Piraí, tributary to rio Negro between Bagé and Dom Pedrito, Bagé. Rio Grande do Sul State. Jacuí basin: MCP 30714, 1, 230.7 mm SL, rio Jacuí-Mirim, Carazinho; MZUSP 25026, 2, 215.9- 225.4 mm SL, rio Jacuí at Ernestina dam near Passo Fundo. URUGUAY. Artigas. Uruguay basin: MCP 17616, 1, 79.2 mm SL, arroyo Arumbeba, circa of 22 km Southeast of Artigas; UFRGS 8059, 1, 101.3 mm SL, arroyo Mandiyú, tributary to río Uruguay, Artigas. Cerro Largo. Uruguay basin: MCP 9878, 1, 288.6 mm SL; MCP 9880, 1, 301.9 mm SL; MCP 9896, 1, 506.0 mm SL; MCP 9926, 1, 399.7 mm SL, río Negro at Arreria. Tacuarembó. Uruguay basin: MZUSP 52006, 4, 289.6- 321.9 mm SL, río Negro, Rincón del Bonete.

Diagnosis. Hoplias lacerdae differs from the other species of H. lacerdae group in the number of pores of the laterosensory system along the ventral surface of dentary (6- 8 vs. 4-6 in H. brasiliensis and H. intermedius , always 5 in H. australis and 4 in H. curupira ). It can be further distinguished from H. brasiliensis and H. curupira , in the number of scales along lateral line (43-48 vs. 38-43 and 34-39 respectively).

Description. Morphometric data presented in Table 7. Body cylindrical, deeper than wide. Greatest body depth at vertical through fifth scale anterior to dorsal-fin origin in specimens smaller than 70 mm SL, closer to dorsal-fin origin in larger specimens. Anterior profile of head slightly rounded in lateral view, more angular in specimens smaller than 160 mm SL. Dorsal profile of head almost straight. Dorsal margin of orbit at horizontal through of dorsal profile of head in specimens smaller than 100 mm SL but not reaching dorsal profile of head in larger specimens. Dorsal profile of body slightly convex from vertical through first series of scale of body to dorsal-fin origin; straight and posteroventrally inclined along dorsal-fin base; straight and less inclined to slightly concave from vertical through base of last dorsal-fin ray to origin of dorsal most procurrent caudal-fin ray. Latter portion slightly more concave in specimens smaller than 100 mm SL. Ventral profile of lower jaw distinctly angular in region of mandibular symphysis, straight to slightly inclined from vertical through anterior nostril to posterior margin of lower jaw. Medial margins of contralateral dentaries running in parallel ( Fig. 1 ac). Ventral profile of body slightly convex to pelvic-fin origin; straight to approximately convex from latter point to anal-fin origin; straight and posterodorsally inclined along anal-fin base; straight to slightly concave from last anal-fin ray to anterior most ventral procurrent caudal-fin ray.

Upper jaw slightly shorter than lower jaw, more so in larger specimens (ca. 380 mm SL). Posterior portion of maxilla dorsally enlarged and extending medially to anterior margins of second and third infraorbitals. Upper and lower lips fleshy with short skin projections covering canines externally. Anterior nostril tubular with anterior slit along its distal half. Anterior and posterior nostrils situated along horizontal through center of orbit, anterior nostril located at approximate one orbital diameter from anterior margin of orbit; posterior nostril midway between anterior nostril and anterior margin of orbit. Eye proportionately larger in smaller specimens. Infraorbital bones well developed and horizontally elongate. Preopercle bearing inumerous laterosensory system pores and partially covered by posterior margin of infraorbitals 3, 4, and posteroventral portion of infraorbital 5. Anteroventral margin of infraorbital 3 relatively straight and posteroventral margin convex. Posterior margin of infraorbital 4 relatively straight and margins of infraorbitals 5 and 6 slightly convex. Small specimens (ca. 70 mm SL) with infraorbital 3 barely reaching orbital rim and proximal ends of infraorbitals 2 and 4 in contact. Larger specimens (ca. 160 mm SL) with infraorbital 4 barely reaching orbital rim and proximal ends of infraorbitals 2 and 5 almost in contact. Specimens larger than 380 mm SL with infraorbital 5 progressively separated from rim. Proximal ends of infraorbitals 2 and 6 almost meeting in some of largest specimens examined (ca. 750 mm SL).

Teeth in both jaws conical or canine. Premaxillary teeth in single row. First premaxillary tooth large canine, and fifth tooth medium sized. Sixth tooth canine and almost as large as anterior most premaxillary canine. Second to fourth, seventh and eighth premaxillary teeth small. Maxilla with single row of approximately 38 relatively small teeth, except for very well developed canine-like fourth tooth. Dentary with anterior external row of teeth and posterior internal row. External series with three anterior small teeth, followed by two well developed canines, with posterior canine larger than anterior canine and then ten conical teeth, slightly smaller than anterior most dentary canine. Internal series beginning at level of last conical tooth of external row and composed of approximately 15 very small teeth. Accessory ectopterygoid and ectopterygoid toothed. Ectopterygoid with series of small conical teeth along its ventrolateral margin and many smaller viliform teeth medially on its ventral surface. Endopterygoid edentulous.

Distal margins of all fins rounded. Total dorsal-fin rays 12-15 (n = 90; ii,10-13; mode: ii,11, n = 66). Dorsal fin located at midbody, its origin at vertical through approximately fourth scale anterior on series along pelvic-fin origin. Longest dorsalfin ray approximately three-quarters of body depth in specimens smaller than 160 mm SL and almost half of body depth in larger specimens. Anal-fin base short. Total analfin rays 9-12 (n = 89; ii,7-10, mode: ii,8, n = 56). Tip of depressed dorsal fin reaching vertical through anal-fin origin in smaller specimens (<160 mm SL), but falling short of vertical through that point in larger specimens. Total pectoralfin rays 11-15 (n = 89; i,10-14; mode: i,12, n = 35). Pectoral-fin origin located at vertical through central portion of opercle. Tip of pectoral fin separated from pelvic-fin origin by five to nine scales. Pectoral and pelvic fins of similar size, slightly larger than anal fin in specimens larger than 380 mm SL. Pelvic-fin rays i,7 (n = 86). Pelvic-fin origin situated at midbody approximately four scales posterior to vertical through dorsal-fin origin. Tips of pelvic fin separated from vertical through anal-fin origin by five to seven scales. Caudal-fin rays i,15,i (n = 15).

Well developed cycloid scales imbricated along body. Dorsal scales begin in series at posterior margin of parietals and overlap supraoccipital spine. Last vertical series of scales on caudal peduncle forms almost straight line on caudal-fin base in specimens smaller than 160 mm SL, and slightly convex arch on caudal-fin in larger specimens. Anterior margin of scales with small recess and posterior margin rounded. Eight to eleven radii extending from center of scale to its anterior margin and around eighteen radii, some anastomosed, extending from center of scale to its dorsal, posterior and ventral margins. Lateral line straight and complete, extending from posteroventral margin of supracleithrum to posterior most scale in body. Lateral line with 43-48 (n = 89, 43 in one specimen; mode: 46, n = 28) perforated scales ( Table 2). Lateral-line scales with single laterosensory canal. Longitudinal series of scales between lateral line and dorsal-fin origin 4.5-6.5 (n = 88; mode: 5.5, n = 61). Longitudinal series of scales between lateral line and pelvic-fin origin 4.5-6.5 (n = 86; mode: 4.5, n = 63). Longitudinal series of scales around caudal peduncle 18-22 (n = 88; mode: 20, n = 73).

Approximately 11 gill rakers on first epibranchial, most in form of small denticulated plates. Lower branch of first branchial arch with five to seven more elongate rakers and approximately 12 plate-like rakers (n = 18). Laterosensory canal along ventral surface of dentary with 6-8 pores (n = 90; mode: 7, n = 67) ( Table 3).

Color in alcohol. Ground coloration of head and body dark to light brown, darker dorsally and paler ventrally. Ventral region white to light yellow in most specimens. Single dark midlateral stripe along lateral line stripe more evident in specimens smaller than 100 mm SL. Midlateral stripe crossed by five to seven dark bars, each three to five scales wide. Diagonal bars in many specimens continuing posteroventrally approximately two or three longitudinal series of scales ventral of lateral line, resulting in chevron-like pattern more conspicuous in specimens smaller than 160 mm SL. Dorsal portions of bars more evident than ventral region. Ground coloration of dorsal surface of head, infraorbital region and opercular series dark brown, with dark reticulated pattern on dorsal surface of head in some specimens. Lips with alternating dark and light vertical bands in specimens smaller than 160 mm SL and completely dark in larger specimens. Coloration of infraorbital region similar to dorsal surface of head. Many specimens with one or two dark stripes extending radially and posteriorly from eye across opercular series and third stripe extending posteroventrally from eye across posterior region of maxilla. Ground coloration of opercular series dark brown. Opercular membrane usually lighter than opercle.

All fins light brown (lighter than body) with dark spots on rays and interradial membranes forming pattern of irregular dark stripes. Stripes of dorsal and caudal fins wider than those of anal fin. Stripes less regular in dorsal fin. Ventral surfaces of pectoral and pelvic fins either lighter than or with same pattern as dorsal surface, but with pattern less conspicuous.

Distribution. Known from the type-locality and its vicinity, rio Ribeira de Iguape basin, and from the rio Uruguay basin, including the río Negro in Uruguay ( Fig. 3 ).

Remarks. Hoplias lacerdae has been regarded as having a wide distribution, occurring in the rio Ribeira de Iguape as well as in rivers of the Amazon, Paraná, São Francisco, Uruguay and other basins. Results presented herein, however, demonstrate that specimens found in the rio Ribeira are morphologically different from those of remaining cited basins with the exception of rio Uruguay. The distribution of Hoplias lacerdae is therefore restricted to rio Ribeira de Iguape and rio Uruguay, being in sympatry with Hoplias australis in the latter basin.

Although samples from the rio Ribeira and rio Uruguay did not differ significantly in the meristic and morphometric features examined in the present study, some geographical disparities should be pointed out. Most specimens from the rio Ribeira have 6.5 and 5.5 scales above and below lateral line, respectively, while the majority of specimens from the rio Uruguay have 5.5 and 4.5 scales above and below lateral line respectively. Specimens from the rio Ribeira tend to have fewer lateral line scales than do specimens from the rio Uruguay (modes: 44 and 46, respectively) and more pectoral-fin rays than do samples from the rio Uruguay (i,13-14 vs. i,10-13, respectively). Nevertheless, these subtle differences must be qualified by the fact that they are not absolute and specimens of H. lacerdae from the rio Ribeira are rare in collections (only ten specimens were available for this study). More specimens from rio Ribeira and the use of cytogenetics and molecular approaches may determine whether these two populations constitute different species.

Bertolletti et al. (1990) and Pessano et al. (2005) cited H. lacerdae from the rio Uruguay basin, the former in the area of Garabi Hydroelectric Station and the latter in Arroio Quarai- Chico. These authors, however, did not mention the basis of those identifications, nor did they illustrate their material. Therefore, it is impossible to determine whether they were dealing with Hoplias lacerdae or the new species, H. australis , described herein, both of which occur in the rio Uruguay basin.

A few specimens of Hoplias lacerdae from Ernestina reservoir in the rio Jacuí, a river running into the Atlantic Ocean in Rio Grande do Sul, southern Brazil, were examined. Malabarba (1989: 127) and Carlos Lucena (pers. comm.) mentioned that the species was introduced from rio Uruguay to rio Jacuí system in the 1980s. Native occurrence of Hoplias lacerdae is therefore considered restricted to the rio Uruguay and rio Ribeira de Iguape drainages. Zaniboni Filho et al. (2004) reported some biological features of this species in the rio Uruguay including length, weight, and growth curve. According to them, the species has fractional spawning and an economic importance in local fisheries and aquacultures.

In the rio Ribeira de Iguape, the species is regarded as rare by local fisherman and a two-year survey of its ichthyofauna failed to yield specimens of this species ( Oyakawa et al., 2006). The small number of specimens of Hoplias lacerdae from rio Ribeira in ichthyological collections may be additional evidence that this species is uncommon, requiring further studies on its ecology and biology and special attention to its conservation status.

Figures

Fig. 10. Hoplias lacerdae, MZUSP 61748, 150.5 mm SL, rio Ribeira de Iguape, São Paulo State, Brazil.

Fig. 11. Hoplias lacerdae, holotype, MNRJ 211, 750.0 mm SL, rio Ribeira de Iguape, São Paulo State, Brazil.

Fig. 1. Ventral view of head of (a) Hoplias curupira, MZUSP 24403, 55.5 mm SL; (b) H. curupira, MZUSP 24403, 92.2 mm SL; (c) H. curupira, MZUSP 24403, 162.2 mm SL and (d) H. malabaricus, MZUSP 90416, 101.7 mm SL.

Fig. 3. Map of South America showing geographic distribution of species of the Hoplias lacerdae group based on material examined in this study. Some symbols may represent more than one lot. Numbers indicate type localities of (1) Hoplias australis (squares); (2) H. brasiliensis (losanges); (3) H. curupira (triangles); (4) H. intermedius (dots); and (5) H. lacerdae (inverted triangles).