A new Hasemania Ellis from the upper rio Paraná basin , with the redescription of Hasemania crenuchoides Zarske & Géry ( Characiformes : Characidae )

A new Hasemania species is described from the headwaters of the rio Uberaba, rio Grande basin, State of Minas Gerais, Brazil. The new species is distinguished from its congeners by a combination of a black, vertically-elongate humeral spot, a single ossification in the position primitively occupied by infraorbitals four and five, four teeth on the inner series of premaxilla, and scales covering the anal-fin base. Hasemania crenuchoides is redescribed and its known geographic distribution is extended; it can be differentiated from other species of the genus by the combination of a black verticallyelongate humeral spot, ii,8 dorsal-fin rays, 11-14 branched anal-fin rays, the presence of separate infraorbitals four and five, and the presence of scales covering the anal-fin base. Phylogenetic relationships of H. crenuchoides and the new species within Hasemania are discussed.


Introduction
Hasemania Ellis is a small genus of the Characidae restricted to the rivers of the Crystaline Brazilian Shield.The genus was proposed by Ellis (1911) to include three species, H. melanura Ellis (type species), H. maxillaris Ellis and H. bilineata Ellis, and differed from all other characids primarily in the absence of the adipose fin.Ellis (1911) further defined the genus by having two rows of premaxillary teeth, few or no teeth along the upper angle of the maxillary, an incomplete lateral line, the caudal fin naked, and a frequently larval pectoral fin in small specimens.Hasemania now includes eight species: H. melanura and H. maxillaris, from the rio Iguaçu basin, H. nana (Lütken) from the rio São Francisco basin, H. hanseni (Fowler) from Goiás (precise locality unknown), H. crenuchoides Zarske & Géry, from the upper rio Paraná basin, H. nambiquara Bertaco & Malabarba, from the upper rio Tapajós basin, H. piatan Zanata & Serra, from the rio de Contas basin, and H. kalunga Bertaco & Carvalho, from the upper rio Tocantins.
Hasemania crenuchoides was until recently only known from its type specimens, and these specimens exhibit a very clear brown staining, without any spot in the body or fins.Zarske & Géry (1999), in the original description of this species, mentioned the specimens preserved for more than thirty years, having a uniform yellowish to olive green coloration, without spots.Recently, Langeani et al. (2007), cited this species from headwaters of the rio Uberaba, rio Grande basin, State of Minas Gerais.Numerous other specimens were subsequently collected in the córrego Paranoazinho, rio Paranaíba drainage, near the type locality of the species; these specimens showing a conspicuous color pattern, including a black humeral spot.
The analysis of this material revealed that the specimens from the rio Paranaíba basin are indeed H. crenuchoides, whereas samples from the rio Grande basin represents a new species, which is described herein together with a redescription of H. crenuchoides.

Material and Methods
Counts and measurements follow Fink & Weitzman (1974).Vertebrae, branchiostegal rays, gill-rakers, total dentary teeth counts and teeth cusp numbers were taken from cleared and stained specimens (c&s) prepared according to Taylor & Van Dyke (1985).All measurements were taken point-to-point with an electronic caliper on the left side of specimens.Standard length (SL) is presented in mm, and the other measurements are expressed as percentages of the SL, except for subunits of head which are presented as percentages of the head length (HL).Vertebral count includes the Weberian apparatus as four elements and the fused PU1+U1 as a single element.Meristic data are presented in the description and are followed by the frequency for each count in parenthesis.An asterisk indicates counts of the holotype.Larger specimens were dissected to observe the gonads for sex determination. Examined

Color in alcohol.
Preserved specimens with dark bodies, chromatophores concentrated mainly dorsally and along posterior margin of scales, resulting in slightly reticulated pattern.Irregular, vertically elongate humeral spot.Longitudinal, mid-lateral stripe extending from vertical midway between head and dorsal-fin origin posteriorly to middle caudal-fin rays.Fins with chromatophores more concentrated along rays than on membranes (Figs.1-2).
Color in life.Live specimens have conspicuous reticulated color pattern, scales with metallic bluish-purple reflections.Dorsal, anal, and caudal fins, mainly the last one, reddish with dark rays.
Sexual dimorphism.Females with a conspicuous concavity along dorsal profile between vertical passing through the posterior nostril to the posterior portion of the head (Fig. 2).Bony hooks along pelvic and anal-fin rays were not observed in any specimen.Etymology.The species name, uberaba, is in reference to the rio Uberaba, where the species is found.A noun in apposition.
Conservation status.Hasemania uberaba until now was only confirmed to occur in the type-locality (area of occupancy AOO less than 10 km2).There is a putative new reference for the species, based on a fish drawing (fig. 2A, Uieda et al., 1987, referred as Hasemania melanura), in the rio Claro, rio Paranaíba basin, a locality very near the type-locality; this additional occurrence, however, could not be confirmed because the specimens analyzed by Uieda et al. (1987) were not conserved and all recent efforts to collect the species in the rio Claro were unsuccessful.The small geographic area (one location) harbors a very peculiar kind of habitat (see "Ecological notes" above) and is limited by a paved road (BR 262) and land areas with agriculture.Furthermore, the waters of the rio Uberaba are used by human consumption and since the southeastern Brazil is suffering a long drought (Escobar, 2015), this can contribute to a decay of the water levels in the basin, which now seems to be a serious plausible threat that can drive the species to critically endangered or extinct in a very short time, allowing to its classification as Vulnerable (VU D2) according to the IUCN (2001IUCN ( , 2011)).In spite of that, it is now preferable to consider Hasemania uberaba as Data Deficient (DD) until more information about its biology and geographical distribution is available, as already pointed out by Carvalho & Langeani (2013) for Hyphessobrycon uaiso, a syntopic characid species.
Color in alcohol.Based on recently collected specimens.Preserved specimens with yellowish bodies and darkbrown chromatophores concentrated more dorsally and along posterior margin of scales (mainly in midportion), resulting in slightly reticulated pattern on body; humeral spot irregular, inconspicuous and vertically elongated, midlateral stripe extending from slightly behind humeral spot to middle caudal-fin rays.Fins with chromatophores more concentrated along rays than on membranes .
Color in life.Body and head yellowish, more so along ventral portion of head, opercular series and anterior portion of body.Pectoral fin yellowish.Dorsal fin mostly hyaline.Pelvic fin reddish to yellowish.Anal fin red overall but turning yellowish along posteriormost rays.Caudal fin reddish along most superior and inferior rays and yellowish along median ones.
Sexual dimorphism.No secondary sexual characters were observed.
Distribution.Hasemania crenuchoides seems to be restricted to the streams from the upper rio Paraná and upper rio Tocantins in the Distrito Federal, Brazil.This is the first reference to the species for the rio Tocantins basin.

Discussion
The relationships of Hasemania uberaba and H. crenuchoides within the genus remain to be investigated.Nevertheless, the two species are probably closely related to H. piatan and H. kalunga, by sharing an apparently derived character, the presence of a rhinosphenoid entirely cartilaginous (vs.ossified).In the other species of Hasemania and small characids the rhinosphenoid is completely ossified, whereas in H. uberaba, H. crenuchoides, H. piatan, and H. kalunga a cartilage occurs in the position primitively occupied by the bone.Additionally, H. uberaba, H. kalunga and H. piatan share the presence of a single ossification in the position primitively occupied by infraorbitals four and five.Hasemania uberaba and H. piatan, in turn, share 18 principal caudal-fin rays (vs.19 in other congeners, and most other characids and characiforms).Zanata & Serra (2010) have already proposed H. piatan as probably phylogenetically close to H. crenuchoides based in the presence of five infraorbitals and a cartilaginous rhinosphenoid.In spite of that, H. crenuchoides, H. uberaba, H. kalunga and H. piatan seem indeed to be closely related based on a combination of cartilaginous rhinosphenoid, larger and deeper bodies, and similar color pattern, with an irregular and vertically elongated humeral spot (vs.rhinosphenoid osseous, body shallow and small and humeral spot absent or rounded and horizontally elongated).

Fig. 3 .
Fig. 3. Jaws of Hasemania uberaba, left side, paratype, DZSJRP 8730, 36.8 mm SL.Scale bar = 1.5 mm.Distribution.Hasemania uberaba is only known from its type locality in the headwaters of the rio Uberaba, rio Grande basin, upper rio Paraná basin, Minas Gerais, Brazil (see Carvalho & Langeani, 2013: figs.4-5).Ecological notes.Headwaters of the rio Uberaba are a marshland with abundant Cyperacea, Poacea and algae, crystalline water and muddy bottom(Carvalho & Langeani,  2013: fig.5).Hasemania uberaba was captured jointly with a new genus and new species of Crenuchidae, the most abundant species (under analysis), Hyphessobrycon uaiso Carvalho & Langeani, and Rivulus sp.The larger, less numerous, specimens among H. uberaba could be easily seen among the smaller specimens of the other species.They swim in open water or sheltered areas with submerged vegetation.Among the specimens examined for stomach contents, one consumed angiosperms and Chironomidae