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Zootaxa 1386: 59 68 (2007) www.mapress.com/zootaxa/ Copyright 2007 Magnolia Press ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) Corumbataia britskii (Siluriformes: Loricariidae: Hypoptopomatinae) a new species from the upper Rio Paraná basin, Mato Grosso do Sul, Central Brazil KATIANE M. FERREIRA 1 & ALEXANDRE C. RIBEIRO 1,2 1 Laboratório de Ictiologia de Ribeirão Preto (LIRP), Departamento de Biologia, FFCLRP-USP, Av. Bandeirantes, 3900, 14040-901, Ribeirão Preto, SP, Brazil. E-mail: katiane@usp.br 2 Departamento de Zoologia, Instituto de Biociências, Universidade Estadual Paulista Júlio de Mesquita Filho (UNESP-Botucatu), Caixa Postal 510, Botucatu, SP, Brazil. E-mail: acribeiro@click21.com.br Abstract A new Hypoptopomatinae, Corumbataia britskii, is described based on specimens recently collected in a tributary of the upper Rio Paraná basin, Mato Grosso do Sul State, Brazil. Corumbataia britskii differs from its congeners by having a more depressed head profile (vs. a more rounded head profile in C. cuestae and C. tocantinensis), by having a more developed and conspicuous tuft of enlarged odontodes on the tip of the supraoccipital (vs. its reduced condition found in C. cuestae and C. tocantinensis), and by heaving a partially enclosed arrector fossae (vs. an almost completely enclosed arrector fossae in C. cuestae and C. tocantinensis). Key words. Siluriformes, Taxonomy, Upper Rio Paraná, South America, Neotropical Region Introduction The genus Corumbataia belongs to the Hypoptopomatinae, a group of small-sized armored catfishes composed by 17 genera and approximately 80 species (Schaefer, 2003; Ribeiro et al. 2005) distributed throughout cis-andean South America from Venezuela to Northern Argentina (Schaefer, 1998). Corumbataia was erected by Britski (1997) who defined the genus by the following combination of characters: exposed portion in the middle of the scapular bridge formed only by the coracoid (i.e., narrow cleithra which are covered by skin medially), compound hypurals 1 and 2 completely fused to the compound hypurals 3 5, atrophied maxillary barbel, and an infraorbital canal entering the infraorbital series via the pterotic-supracleithrum. Corumbataia cuestae occurs in small streams of the Rio Tietê (upper Paraná basin), and C. tocantinensis occurs in the Rio Araguaia and Rio Tocantins. In this paper, we describe a third species of Corumbataia, from a headwater stream of the Rio Sucuriú, upper Paraná River Basin, Mato Grosso do Sul State, Central Brazil. Material and methods Measurements follow Boeseman (1968), including anal width, folded dorsal-fin length, snout-opercle length (Armbruster & Page, 1996) and preanal length (Ribeiro et al. 2005). Plate counts and nomenclature follow the schemes of serial homology proposed by Schaefer (1997). Meristic data follow Bockmann & Ribeiro (2003). Vertebral counts included all vertebrae (including the first five vertebrae modified into Weberian Apparatus), with the compound caudal centrum (PU1+U1) (Lundberg & Baskin, 1969) counted as a single element. All Accepted by M. de Carvalho: 17 Nov 2006; published: 4 Jan. 2007 59

morphometric and meristic data, including premaxillary and dentary teeth counts, were taken from the left side of the specimens. Osteological terminology follows Schaefer (1997). Osteological examination and plate counts were performed on specimens cleared and stained (c&s) according to the procedure of Taylor & Van Dyke (1985). Institutional abbreviations are as follows: LIRP (Laboratório de Ictiologia de Ribeirão Preto, Departamento de Biologia da FFCLRP, Universidade de São Paulo, Ribeirão Preto, Brazil) and LBP (Laboratório de Biologia de Peixes, Departamento de Morfologia, Instituto de Biociências-UNESP, Botucatu, Brazil). Corumbataia britskii, new species Fig. 1 Holotype. LIRP 5884, 26.0 mm SL, Brazil, State of Mato Grosso do Sul, Chapadão do Sul, Córrego Pedra Azul (19º12 15 S 52º43 55 W), Rio Sucuriú, upper Rio Paraná basin (Fig. 2), 10 October 2003, K.M. Ferreira, R.Campos-da-Paz & E. Sá. FIGURE 1. Lateral, dorsal and ventral views of holotype of Corumbataia britskii, LIRP 5884, 26.0 mm SL. Paratypes. LIRP 4937, 5, 13.3 27.0 mm SL (2, 19.7 24.4 mm SL, c&s), as the holotype. All from Brazil, Mato Grosso do Sul State, Chapadão do Sul, Rio Sucuriú, upper Rio Paraná basin: LIRP 5897 (2, 17.0 23.6 mm SL, c&s), Córrego Pedra Azul (19º12 15 S 52º43 55 W), 23 August 2006, A.C. Ribeiro, M. Carvalho & 60 Zootaxa 1386 2007 Magnolia Press FERREIRA & RIBEIRO

O. Froehlich; LIRP 5898, (8, 17.6 24.7 mm SL), Córrego Fazendinha (19º14 43 S 52º39 11.2 W), 25 August 2006, A.C. Ribeiro, M. Carvalho & O. Froehlich. FIGURE 2. Type locality of Corumbataia britskii. Córrego Pedra Azul (19º12 15 S 52º43 55 W), Rio Sucuriú, upper Rio Paraná basin, Central Brazil. Diagnosis. Corumbataia britskii differs from congeners by having a more depressed head profile (vs. more rounded in C. cuestae and C. tocantinensis) (Fig. 3), by having a more developed and conspicuous tuft of enlarged odontodes on the tip of the supraoccipital (vs. its reduced condition found in C. cuestae and C. tocantinensis) (Fig. 3), and by heaving a partially enclosed arrector fossae (vs. an almost completely enclosed arrector fossae in C. cuestae and C. tocantinensis) (Fig. 4). Description. Morphometric and meristic data given in Tables 1 and 2, respectively. Dorsal profile of the body convex from snout tip to dorsal-fin origin; slightly concave at dorsal-fin base; straight from end of dorsal-fin base to caudal-fin origin. Ventral profile of the body straight from the snout tip to anus; straight at analfin base; straight from end of anal-fin base to caudal-fin origin. Greatest body depth at dorsal-fin origin; least body depth at caudal peduncle. Greatest body width at opercular or cleithral regions. Body progressively narrowing caudally from cleithrum. Head rounded anteriorly in dorsal view; eyes dorsolaterally positioned. Oral disk slightly elliptical, narrowing posteriorly. Anterior oral disk lobe covered by papilla of about same size. Posterior disk lobe covered by papilla which decrease in size posteriorly. Maxillary barbel atrophied. Compound pterotic-supracleithrum-posttemporal with small perforations along its median and inferior margins. Body entirely covered by plates, except on rostrum, region overlying opening of swim bladder capsule, around anus, and in pectoral-, pelvic- and anal-fin insertions. Scapular bridge exposed and covered by odontodes. Abdomen in adult specimens covered with small plates which are absent in young specimens. Body plates completely covered by odontodes. Larger odontodes neighboring the supraoccipital bone. Lateral line interrupted by gaps in middle region and in caudal peduncle in a young c&s specimen, but not interrupted in larger specimens. Dorsal-fin origin located posteriorly to vertical through pelvic-fin origin. Adpressed pectoral fin extending over the pelvic-fin origin. Tip of pelvic fin reaching anal-fin origin in males and shorter in females. Color in alcohol. Dark brown coloration on dorsal and lateral regions of body; ventral region yellowish and belly slight grayish. Numerous dark brown chromatophores distributed over head, forming a diffuse stripe from snout tip below eyes to posterior region of head. Chromatophores concentrated on dorsal- and anal-fin bases. Dark brown stripe extending from end of dorsal- to caudal-fin origin. Large diffuse dark brown midlat- A NEW CORUMBATAIA (SILURIFORMES) Zootaxa 1386 2007 Magnolia Press 61

eral stripe from snout tip, crossing eyes, extending to caudal-fin base. Large ventral dark brown stripe from around urogenital papillae to caudal-fin origin. Abdomen mostly unpigmented. Three to four transverse darkbrown stripes on dorsal-fin. Unbranched rays of pectoral, pelvic and anal fins with several scattered, widespread chromatophores not arranged in stripes. Caudal-fin mostly dark-brown with two unpigmented round spots on caudal-fin lobes. TABLE 1. Descriptive morphometrics of Corumbataia britskii. Values are given for the holotype (separately) and 14 paratypes. Hol. = Holotype; SD = Standard Deviation. Character Hol. Minimum Maximum Mean SD Standard length (mm) 26.0 13.3 27.0 21.4 3.4 Percents of standard length Predorsal length 42.7 42.5 48.3 45.5 2.0 Preanal length 61.2 48.3 63.4 54.8 5.0 Head length 31.9 31.5 36.6 34.1 1.9 Cleithral width 25.4 18.0 25.9 21.9 3.4 Dorsal-fin spine length 22.3 17.6 23.6 20.8 1.9 Folded dorsal-fin length 25.8 18.4 26.9 24.2 2.2 Base of dorsal-fin length 15.4 11.3 15.4 13.5 1.1 Thorax length 21.2 16.0 21.2 18.3 1.3 Pectoral-fin spine length 21.9 16.8 23.6 19.5 2.5 Abdomen length 18.8 11.3 22.7 16.4 4.3 Pelvic-fin spine length 18.1 15.5 21.4 18.4 2.2 Postanal length 28.5 24.4 32.1 28.3 2.4 Caudal-peduncle depth 10.4 8.3 10.8 10.2 0.7 Anal width 6.9 5.7 8.9 7.3 0.8 Snout-opercle length 24.2 20.9 26.4 24.4 1.3 Percents of head length Head width 76.6 63.9 78.4 72.8 4.0 Head depth 99.5 92.5 107.2 100 5.2 Snout length 39.5 34.8 44.9 38.0 2.7 Interorbital length 43.3 32.8 43.3 38.9 2.9 Orbital diameter 14.3 13.5 19.8 16.7 2.2 Percents of cleithral width Anal width 50.8 34.1 52.1 43.8 5.8 Ecological notes. The new species was sampled in deforested areas in moderate to fast current streams. Corumbataia britskii was sampled in association with aquatic macrophytes or in the submerged portion of marginal vegetation. In its gut contents were found filamentous blue-green algae, chlorophyts, diatoms and periderm. Distribution. Known just from two streams, Córrego Pedra Azul (type locality) and Córrego Fazendinha (19º12 15 S 52º43 55 W and 19º14 43 S 52º39 11.2 W, respectively). Both streams are tributaries of the Rio Sucuriú, a right tributary of the upper Rio Paraná basin in the Mato Grosso do Sul State, Brazil. 62 Zootaxa 1386 2007 Magnolia Press FERREIRA & RIBEIRO

Etymology. Named after Dr. Heraldo A. Britski, Museu de Zoologia da Universidade de São Paulo (MZUSP), in recognition of his many contributions to our understanding of Hypoptopomatinae catfishes. FIGURE 3. Lateral view of head of Corumbataia. (A) C. britskii, holotype, LIRP 5884, 26.0 mm SL; (B) C. cueatae, LIRP 2774, 29.5mm SL; (C) C. tocantinensis, LBP 1653, 26mm SL. Arrows indicate tip of supraoccipital. A NEW CORUMBATAIA (SILURIFORMES) Zootaxa 1386 2007 Magnolia Press 63

TABLE 2. Frequency distribution and summary of meristics for Corumbataia britskii. Holotype values are marked with an asterisk. Meristics marked with double asterisks were counted in c&s specimens only. Character Frequency Distribution Range Mode Dorsal plates 22 (2); 23 (4); 24 (1) 22 24 23 Mid-dorsal plates 18 (3); 19 (1) 18 19 18 Median plates 19 (3); 20 (5); 21 (4) 19 21 20 anterior field plates 8 (1) --- --- gap field plates 6 (1) --- --- posterior field plates 6 (1) --- --- Mid-ventral plates 19 (1); 20 (2) 19 20 20 Ventral plates 18 (1); 19 (1); 20 (4) 18 20 20 Predorsal plates 3 (16) --- --- Dorsal plates below dorsal-fin base 5 (16); 6* (2) 5-6 5 Dorsal plates between end of dorsal base and caudal-fin membrane 12 (16) --- --- Ventral plates above anal-fin base 3 (12); 4 (3) 3 4 3 Ventral plates between end of anal base and caudal-fin membrane 9* (14); 10 (1) 9 10 9 Dorsal-fin branched rays 7 (all specimens) --- --- Pectoral-fin branched rays 6 (all specimens) --- --- Pelvic-fin branched rays 5 (all specimens) --- --- Anal-fin branched rays 5 (all specimens) --- --- Caudal-fin branched rays 14 (all specimens) --- --- Dorsal procurrent rays ** 3 (all specimens) --- --- Ventral procurrent rays ** 3 (all specimens) --- --- Total vertebrae ** 27(1); 28(1) 27 28 --- Premaxillary teeth 27 (3); 28* (5); 29 (4) 27 29 28 Dentary teeth 23* (4); 24 (3); 25 (2); 26 (3); 27 (1) 23 27 23 Discussion The phylogenetic relationships of Corumbataia is presently somewhat obscure. Schaefer (1991, 1997, 1998) provided a phylogenetic analysis of the Hypoptopomatinae at the level of genera, recognizing it as a monophyletic unit and providing evidence for two natural subgroups within the subfamily, the tribes Otothyrini and Hypoptopomatini. Britski (1997), based on the set of characters presented by Schaefer (1991), included Corumbataia in the Otothyrini. This generic placement was not tested subsequently by Schaefer (1998), who did not include Corumbataia in his analysis. Recent papers (Ribeiro et al. 2002; Ribeiro et al. 2005) concerning the inclusion of new taxa within Schaefer s phylogenetic framework have not provided substantial changes in hypoptopomatine topology, and are in agreement with Schaefer s main conclusions. However, Gauger & Buckup (2005), in including in a phylogenetic analysis two new species of Parotocinclus, several 64 Zootaxa 1386 2007 Magnolia Press FERREIRA & RIBEIRO

undescribed cascudinhos, as well as Corumbataia, concluded that Schaefer s Otothyrini is not monophyletic. These authors presented two alternative cladograms in their reanalysis of the Hypoptopomatine. In one of them, Corumbataia appears in a group that includes Microlepidogaster, Pseudotocinclus, Schizolecis, Pseudotothyris and Otothyris, all of which belonged to Schaefer s original concept of Otothyrini, corroborating the previous generic placement of Corumbataia made by Britski (1997). FIGURE 4. Ventral view of pectoral girdle of species of Corumbataia. (A) C. britskii; (B) C. cuestae; (C) C. tocantinensis. A NEW CORUMBATAIA (SILURIFORMES) Zootaxa 1386 2007 Magnolia Press 65

FIGURE 5. Lateral view of the caudal-fin indicating color patters in Corumbataia. (A) female of C. britskii; (B) male of C. britskii (arrows indicate clear caudal-fin blotches of males); (C) C. tocantinensis; (D) C. cuestae (arrows indicate stripes on unbranched caudal-fin rays). We confirm in C. britskii all of the characters presented by Schaefer (1998) supporting the Otothyrini, except his character 35 (presence of a rostral plate with posterior notch articulating with mesethmoid Corumbataia presents a naked snout tip in which the rostral plate is absent), which are: presence of a deep metapterygoid channel (character 13, state 2), presence of a preopercle with a medially reflected ventral margin (character 18, state1), and presence of a ventrally expanded fourth infraorbital (character 37, state1). According to Britski (1997), C. cuestae and C. tocantinensis should be differentiated only by counts of premaxillary and dentary teeth (23 28, 21 26 vs. 27 40, 26 39, respectively). However, Ferreira et al. (2006) also found citogenetic differences between C. cuestae and C. tocantinensis. Both species present biarmed chromosomes, but in C. cuestae the chromosomes are more metacentric while in C. tocantinensis they are more submetacentric. Furthermore, we observed that the two species can also be differentiated by color pattern, as the unbranched caudal-fin rays of C. britskii and C. tocantinensis lack the striped pattern present in C. custae (Fig. 5). Corumbataia tocantinensis and C. cuestae are more similar to each other in the morphology of the pectoral skeleton (Fig. 4). Both species share almost completely enclosed arrector fossae, which is partially opened in C. britskii. The lateral portions of the cleithra are wider in C. britskii than in the other two species. Britski (1997) mentioned sexually dimorphic features for both C. cuestae and C. tocantinensis. Males differ from females by presenting a developed urogenital papillae posterior to the anus, a skin fold at the dorsal portion of the pelvic-fin spine, and a much longer pelvic-fin spine that extends over the first anal-fin ray. We confirmed all these characters in C. britskii and also observed that males of this species differ from females by having four white blotches on the caudal-fin, two at the dorsal lobe and two at the ventral lobe. Females, on the other hand, present only two white blotches, one on each lobe (Fig. 5A, B). Comparative material. Corumbataia cuestae: MZUSP 51222, Holotype, São Paulo State Analândia, Cachoeira de Analândia, rio Corumbataí, H.A. Bristki, 27/07/1963; LIRP 2774, 169 (6 c&s), Brazil, São 66 Zootaxa 1386 2007 Magnolia Press FERREIRA & RIBEIRO

Paulo State, São Pedro, Ribeirão da Ponta do Meio, Fazenda. Sta. M. da Ponta do Meio (22º33 27 S 47º57 08 W), R.M.C. Castro, F.A. Bockmann, H.F. Santos, T.X. Abreu & G.Z. Pelição, 29/05/2001. Corumbataia tocantinensis: MZUSP 51223, Holotype, Goiás State, rio Vermelho, Expedição DZ (H.A. Britski), 14/ 11/1966; LBP 1653, 27 (4 c&s), Brazil, Goiás State, Rio Araguaia, Rio Vermelho (15º55 1.5 S 50º07 43 W), C. Oliveira, 15/12/2002; LBP 1972, Brazil, Goiás State, Rio Araguaia, Rio Vermelho (15º54 10.9 S 50º06 53.8 W), C. Martins, 30/08/2003. Acknowledgments Murilo Carvalho and Otávio Froehlich helped during a field trip to collect additional specimens of our new species. Murilo Carvalho and Marcelo Britto provided suggestions that improved the manuscript. Cláudio Oliveira (LBP) is thanked for the loan of comparative material. This paper was financially supported by the All Catfishes Species Inventory Project (NSF DEB 0315963) that provided funding to visit collections. We also would like to thank our respective grants for financial support that allowed us to work on this project: Fundação de Amparo a Pesquisa do Estado de São Paulo (FAPESP 02/05464-0 to KMF) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq 140488/04-9 to ACR). References Armbruster, J.W. & Page, L.M. (1996) Redescription of Aphanotorulus (Teleostei: Loricariidae) with description of one new species, A. ammophilus, from the Río Orinoco basin. Copeia, 1996 (2), 379 389. Bockmann F.A & Ribeiro, A.C. (2003) Description of a new suckermouth armored catfish of the genus Pareiorhina (Siluriformes: Loricariidae), from southeastern Brazil. Ichthyological Exploration of Freshwaters, 14 (3), 231 242. Boeseman, M. (1968) The genus Hypostomus Lacépède, 1803, and its Surinam representatives (Siluriformes, Loricariidae). Zoologische Verhandelingen, 99:1 89. Britski, H.A. (1997) Descrição de um novo gênero de Hypoptopomatinae, com duas espécies novas (Siluriformes, Loricariidae). Papéis Avulsos de Zoologia, 40 (15), 231 255. Ferreira, D.C., Chiachio, M.C., Takako, A.K., Andreata, A.A., Foresti, F. & Oliveira, C. (2005) Comparative cytogenetics of nine species of Hypoptopomatinae (Teleostei: Siluriformes: Loricariidae): the importance of structural rearrangements in chromosome evolution. Caryologia, 58(4), 387 395. Gauger, M.F.W. & Buckup, P.A. (2005) Two new species of Hypoptopomatinae from the rio Paraíba do Sul basin, with comments on the monophyly of Parotocinclus and Otothyrini (Siluriformes: Loricariidae). Neotropical Ichthyology, 3(4), 509 518. Lundberg, J.G. & Baskin, J.N. (1969) The caudal skeleton of the catfishes, order Siluriformes. American Museum Novitates, 2398,1 49. Ribeiro, A.C., Melo, A.L.A. & Pereira, E.H.L. (2002) A New Species of Parotocinclus Eigenmann & Eigenmann, 1889 (Siluriformes: Loricariidae) from the Rio São Francisco Basin, Southeastern Brazil. Ichthyological Exploration of Freshwaters, München, 13(3), 217 224. Ribeiro, A.C., Carvalho, M. & Melo, A.L.A. (2005) Description and relationships of Otothyropsis marapoama, a new genus and species of Hypoptopomatinae catfish (Siluriformes: Loricariidae) from rio Tietê basin, southeastern Brazil. Neotropical Ichthyology, 3(4), 489 498. Schaefer, S.A. (1991) Phylogenetic analysis of the loricariid subfamily Hypoptopomatinae (Pisces: Siluroidei: Loricariidae), with comments on generic diagnoses and geographic distribution. Zoological Journal of the Linnaean Society, 102,1 41. Schaefer, S.A. (1997) The Neotropical Cascudinhos: Systematics and biogeography of the Otocinclus catfishes (Siluriformes: Loricariidae). Proceedings of the Academy of Natural Sciences of Philadelphia, 148,1 120. Schaefer, S.A. (1998) Conflict and resolution: Impact of new taxa on phylogenetic studies of the neotropical cascudinhos (Siluriformes: Loricariidae). Pp. 375 400. In: L.R. Malabarba, R.E. Reis, R.P. Vari, C.A.S. Lucena, & Z.M.S. Lucena (Eds.). Phylogeny and Classification of Neotropical Fishes. Porto Alegre, Edipucrs, 603p. Schaefer, S.A. (2003) Loricariidae Hypoptopomatinae (Armored catfishes). Pp. 321 329. In: R.E. Reis, S.O. Kullander & C.J. Ferraris, Jr. (Eds.) Checklist of the Freshwater Fishes of South and Central America. Porto Alegre: Edipucrs, 729p. A NEW CORUMBATAIA (SILURIFORMES) Zootaxa 1386 2007 Magnolia Press 67

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