New species of Hasemania ( Characiformes : Characidae ) from Central Brazil , with comments on the endemism of upper rio Tocantins basin , Goiás State

Hasemania kalunga is described from the upper rio Tocantins basin, Chapada dos Veadeiros, Goiás State, Brazil. The new species is distinguished from its congeners by the number of perforated lateral line scales (11-21 vs. 5-9, except from H. crenuchoides and H. piatan), by the presence of a black vertically-elongate humeral spot (vs. absent, except from H. crenuchoides, H. nambiquara, and H. piatan), and by absence of scale sheath along anal-fin base (vs. presence, except from H. maxillaris, and H. piatan), by 19 principal caudal-fin rays (vs. 18 in H. piatan). It differs also from H. crenuchoides by morphometric data. The absence of scale sheath covering the anal-fin base, an uncommon character in Characidae, and the endemism of the ichthyofauna from the upper rio Tocantins are discussed.


Introduction
Hasemania is a small Characidae incertae sedis genus (sensu Lima et al., 2003), proposed by Ellis (1911) to include H. maxillaris, H. melanura (type species) and H. bilineata (= Coptobrycon bilineatus). It differs from other characids mainly by the absence of an adipose fin. The presence of two series of teeth in the premaxilla, the maxilla with few or no teeth along its exposed anterior margin, the incomplete lateral line, and naked caudal fin led Ellis (1911: 148) to consider the new genus "like Hyphessobrycon, but without an adipose". Later, Eigenmann (1921) recognized the same species of Ellis in his revision. Géry (1977) followed Eigenmann and Ellis, and further included the genus in a group he named "Hemigrammus and allied genera". Publications on Hasemania subsequent to Eigenmann's revision, involve mainly the transfer of Pristicharax hanseni to Hasemania hanseni by Böhlke (1958) and Tetragonopterus nanus to Hasemania nana by Géry (1972), and descriptions of new species by Zarske & Géry (1999), Bertaco & Malabarba (2007), and Zanata & Serra (2010).
Currently, seven valid species are recognized in Hasemania, three with a limited distribution in the rio Paraná basin (H. crenuchoides Zarske & Géry from the upper rio Paraná; H. maxillaris Ellis and H. melanura Ellis from rio Iguaçu), rio São Francisco basin (H. nana); one species from an unknown locality in the state of Goiás, Brazil (H. hanseni (Fowler)), one from the upper rio Tapajós basin (H. nambiquara), and one from the upper rio de Contas drainage (Lima et al., 2003;Bertaco & Malabarba, 2007;Zanata & Serra 2010). In a recent expedition to tributaries from the upper rio Tocantins basin in the Chapada dos Veadeiros, Goiás State, Central Brazil, a new characid species referable to Hasemania was discovered and is described herein. The new species is allocated in Hasemania because it fits best in the traditional diagnosis of the genus given by Ellis (1911) and subsequent authors.

Material and Methods
Counts and measurements follow Fink & Weitzman (1974), with the exception of the number of scale rows below the lateral line, which followed Bertaco & Lucena (2006). Vertebrae, supraneurals, gill-rakers on the first arch, and procurrent caudal-fin ray counts were taken from cleared and stained specimens (c&s) prepared according to Taylor & van Dyke (1985). Vertebral counts include the four vertebrae of the Weberian apparatus, and the terminal centrum counted as a single element. Pattern of circuli and radii was defined on scales sampled from region between the lateral line and the insertion of pelvic-fin. The values for the holotype are indicated by an asterisk. SEM (scanning electron micrographs) photos were taken from the cleared and stained dissected specimen. Measurements were taken point-to-point with an electronic caliper on the left side of specimens. All measurements other than standard length (SL) are expressed as a percentage of SL except for subunits of the head, which are presented as a percentage of head length (HL Description. Morphometric data summarized in Table 1. Body compressed and moderately deep; greatest body depth located between tip of supraoccipital spine and dorsal-fin origin. Dorsal profile of head between vertical through posterior nostril and tip of supraoccipital spine straight or slightly convex. Body profile convex from tip of supraoccipital spine to base of last dorsal-fin ray, and straight from that point to dorsal caudal peduncle margin. Ventral profile of body convex from margin of lower lip to anal-fin origin. Profile of body along anal-fin base posterodorsally slanted. Caudal peduncle deep, profile nearly straight to slightly concave along dorsal and ventral margins. Infraorbital series with five bones. Rhinosphenoid present. Snout rounded from margin of upper lip to vertical through anterior nostrils. Mouth terminal. Maxilla extending posteriorly to vertical through middle of orbit, slightly curved, and oriented at angle of approximately 45 degrees relative to longitudinal body axis. Two tooth rows in premaxilla: outer row with 3-4 (4*) trito pentacuspid teeth, central cusp longer; inner row with 5 teeth, gradually decreasing in length from the first to fourth, last distinctly smaller, with 5-7 cusps; central cusp longer and broader than lateral cusps. Maxilla with 1 to 3 (2*) teeth, each with 3 to 5 cusps with central cusp largest. Four anteriormost dentary teeth largest, with 5-7 cusps, and followed first by somewhat smaller tooth with 3-5 cusps, and then 5-6 distinctly smaller teeth with 1-3 cusps; central cusp in all teeth two or three times longer and broader than remaining cusps. Cusp tips slightly curved posteriorly and lingually (Fig. 2).
Dorsal-fin rays ii,8 (ii,8*, one with ii,7, n = 28); first unbranched ray approximately one-half length of second unbranched ray. Dorsal-fin origin posterior to middle of SL and slightly posterior to vertical through pelvic-fin insertion. Adipose fin absent in all examined specimens. Anal-fin rays ii-iv,13-14 (iii,14*, rarely with 12 or 15, mean = 13.6, n = 28). First unbranched ray only apparent in the cleared and stained specimens. Longest branched anal-fin ray reaching or surpassing origin of last anal-fin ray when fin depressed. Anal-fin origin situated posterior to vertical through base of last dorsal-fin ray. Anal-fin rays of males bearing one pair of small bony hooks along posterolateral border of each segment of lepidotrichia, usually along last unbranched ray to eighth anterior branched rays. Hooks more numerous along second through fifth branched rays, and usually located along posteriormost branch and distal 1 / 2 to 2 / 3 of each ray.
Pectoral-fin rays i,10-13 (11*, mean = 11.2, n = 28). Tip of pectoral fin not reaching pelvic-fin insertion in males and females. Pelvic-fin rays i,6 (n = 28). Tip of pelvic fin reaching anal-fin origin in males, falling short of that point in females. Pelvic-fin insertion located slightly anterior to vertical through dorsal-fin origin. Pelvic fin of males usually bearing one small bony hook per lepidotrichia segment along ventromedial border of first to fifth branched rays.

Color in alcohol.
General body color yellowish to yellowishbrown. Dorsal and dorsolateral portions of head and body pigmented dark brown. Dorsolateral portion of body with scales bordered by dark pigment and forming reticulate pattern. One black humeral spot vertically elongate, wide in dorsal portion and narrowing ventrally; wide dorsal portion of humeral spot located over two or three scales of two to   Color just after fixation. Color pattern similar to described for alcohol preserved specimens. Specimens examined soon after fixation in formalin with all fins orange to yellowish pigmented.
Sexual dimorphism. Males of Hasemania kalunga are easily differentiated by the presence of small bony hooks on the anal-, and pelvic-fin rays (see Description). Also, males and females differ in pectoral and pelvic-fin lengths, body depth (Table 1), and anal-fin shape, which is nearly straight in males and concave in females. Females also attain larger body sizes than males. From 119 specimens fixed in formalin 10% only eight are males (31.0-39.0 mm SL), 26 are females (34.1-53.3 mm SL), and remaining are unsexed specimens (15.5-34.0 mm SL). Gill glands (Burns & Weitzman, 1996) were not found on first gill arch in males and females.
Ecological notes. All specimens of Hasemania kalunga were collected along the margins of semilentic stretches of a shallow, clear water river, greatest depth of 1.20 m, width of 1.5 to 10 m, with sand and leaves on the bottom, with moderate submerged vegetation (mainly Poaceae and Cyperaceae) and a little riparian vegetation (shrubs). Stomach contents of five specimens showed a large volume of Formicidae, followed by Diptera, Clorophyta, unidentified vegetal organic matter, and insects fragments in lesser amount. No other fish species was collected with H. kalunga in the type locality (Fig. 3). The type locality is situated about 8 km upstream from Kalunga community where are located some waterfalls, e.g. Cachoeira da Santa Bárbara and Cachoeira da Capivara.

Distribution.
Hasemania kalunga is known from its type locality in the upper rio Tocantins basin, tributary of rio das Almas, Cavalcante, Goiás State, Brazil (Fig. 4).
Etymology. The species name, kalunga, is in reference to Comunidade Quilombo Kalunga, descendents of African slaves that lives in the region of the upper rio Tocantins basin (Cavalcante, Teresina de Goiás, and Monte Alegre de Goiás municipalities), in the Chapada dos Veadeiros, near the type locality. Kalunga, in Bantu language, mean protected sacred place. A noun in apposition.
Relationships of Hasemania kalunga with other Hasemania species remain to be investigated, but it is close to H. crenuchoides, and H. piatan. The monophyly of Hasemania based solely on the lack of an adipose fin has been questioned by several authors (Böhlke, 1958;Géry, 1972Géry, , 1977Weitzman & Malabarba, 1999), and more recently by Lima & Gerhard (2001), who included a newly described characid species lacking an adipose fin in Hyphessobrycon instead of Hasemania due to an overall body resemblance of the new species, Hyphessobrycon negodagua, with several species of this genus. A similar procedure was adopted previously by Ellis (1911), in the description of Hyphessobrycon taurocephalus that lacks an adipose fin. Hasemania kalunga differs from H. negodagua and H. taurocephalus in its color pattern, in the number of perforated lateral line scales (11-21 vs. 5-11 and 8-10, respectively), and additionally from H. taurocephalus in the number of maxillary teeth (1-3 vs. absent).
Hasemania crenuchoides was described by Zarske & Géry (1999)  Hasemania kalunga is similar to Hyphessobrycon hamatus (upper rio Tocantins basin), but differs in the number of branched anal-fin rays (12-15 vs. 16-18), absence of scale sheath along anal-fin base and of an adipose-fin (vs. presence), and absence of bony hooks in the dorsal and pectoral fin of males (vs. presence) (Bertaco & Malabarba, 2005). Hasemania kalunga is the first species of the genus described from the rio Tocantins basin, possibly due to the limited collecting effort, mainly in the upper portion of this basin. It is one of the largest species of Hasemania along with H. crenuchoides.
The ichthyofauna from the upper rio Tocantins basin, that comprises the drainages above the junction of Paranã and Maranhão or Tocantins rivers (approximately at 12º29'S 48º12'W) (Ribeiro et al., 1995), is greatly endemic. This drainage basin possesses various natural waterfalls along of its rivers or streams that can reach up to 120 meters high. Diverse fish species are endemic from the upper portion of the basin, mainly from its headwaters. Apart from new species, other reports on the species composition of that basin are rare (Miranda & Mazzoni, 2003