A new species of Crenicichla from the upper Rio das Antas basin, dos Patos lagoon system, southern Brazil (Teleostei: Cichlidae)

Crenicichla lucenai sp. n. from the upper Rio das Antas basin, dos Patos lagoon system, southern Brazil is here described. The new species is similar to C. punctata and C. maculata. It is distinguished from both these species by the conspicuous pattern of dark brown irregular lines extending from longitudinal stripe to ventral profile (versus irregular lines absent). In addition, C. lucenai can be distinguished from C. punctata by the absence of dots on the sides of its head.


Introduction
Crenicichla Heckel, 1840 is a Neotropical genus of predatory fish known as pike-cichlids.Currently, this genus comprises more than 85 valid taxa, being the most species rich genus of the family Cichlidae in the Neotropical region (Kullander, 2003;Kullander & lucena, 2006;Montaña et al., 2008;Kullander et al., 2010;PiáleK et al., 2012;Varella et al., 2012;casciotta et al., 2013).The species of Crenicichla occur in the cis-Andean region of South America from the coastal drainages of Venezuela and the Guianas to the lower La Plata river basin in Argentina (Kullander, 2003;casciotta et al., 2006;casciotta et al., 2010;Kullander et al., 2010).Kullander (1998), using morphological characters, considered Crenicichla to be the sister taxon of Cichla Bloch & schneider, 1801, and classified the entity Cich la + Crenicichla as a relatively basal clade of the Neotro pical Cichlidae.However, molecular phylogeny studies strongly support Crenicichla as a member of the derived geophagine cichlids (Farias et al., 2000;lóPez-Fernández et al., 2005a, 2005b, 2010;sMith et al., 2008).Several species groups have been recognized within the genus Crenicichla based on morphological character states (e.g.Kullander, 1991;Ploeg, 1991;lucena & Kullander, 1992;Kullander & lucena, 2006).Recently published molecular phylogenies have questioned the monophyly and validity of some of these species groups, as well as their supposed relationships (Kullander et al., 2010;PiáleK et al., 2012).Kullander & lucena (2006) provided a revision of the Crenicichla species from the coastal river basins of southeastern and southern Brazil.Formerly, these species were pooled in the C. lacustris group sensu Kullander (1982).However, the monophyly and validity of this species group has been put in question (lucena & Kullander, 1992;Kullander & lucena, 2006;Kullander et al., 2010;PiáleK et al., 2012).Following the revision by Kullander & lucena (2006), six species from this geographic region have been recognized as valid: C. iguapina Kullander & lucena, 2006, from the Ribeira do Iguape river drainage; C. tingui Kullander & lucena, 2006, from the Itapocu river drainage and the Babitonga and Paranaguá bays systems; C. maculata Kullander & lucena, 2006, from the coastal rivers' drainages and lake systems lying between north of dos Patos lagoon and the Itapocu river drainage; C. lacustris castelnau, 1855, from Paraíba do Sul and São João river basins and lakes of the upper Doce river drainage; C. punctata hensel, 1870, from the dos Patos lagoon system, including the Mirim lagoon basin; and C. mucuryna ihering, 1914, from the Mucuri river basin (Kullander & lucena, 2006).Kullander & lucena (2006) arranged these species into three distinct morphological groups: C. punctata and C. maculata, characterized by having E1 row scales counts in the range 56 -75, a row of blotches along the side of the body and males being densely spotted on body and fins; C. iguapina, C. tingui and C. lacustris are characterized by having a wide longitudinal stripe along the side, males densely spotted on body and fins, and E1 row scales counts range 60 -75.Crenicichla mucuryna can be distinguished from the previous morphological assemblages by having a trunk with narrow bars and an absence of dots.
The purpose of this paper is to describe a new Crenicichla species originating from the Atlantic coastal river basins of southern Brazil.The new species occurs in the upper Rio das Antas basin, Jacuí river drainage.Based on morphological character states, it is closely related to C. maculata and C. punctata.Kullander &lucena (2006) andottoni et al. (2011).Measurements were taken on the left side of each specimen with digital cali-pers under a binocular microscope.Osteological studies were made on cleared and counterstained (C&S) specimens prepared according to taylor & Van dyKe (1985).Osteological nomenclature follows costa (2006).Nomenclature of the Crenicichla species groups follows PiáleK et al. (2012) and Kullander & lucena (2006).

Measurements and counts follow
For species delimitation we adopted the population aggregation analysis (daVis & nixon, 1992)    terior half of head.Single dorsolateral nostrils (not tubular) between orbit and tip of snout, located at about ¼ of the distance from orbit.Vertical margin of preopercle serrated.
Flank scales strongly ctenoid.All scales on head, anteriorly on dorsum (above anterior ¼ of lateral line), along dorsal fin base, chest, and on ventral parts of body from lower edge of pectoral axilla to anal fin origin, and along anal fin base cycloid.Predorsal scales small and cycloid, covered by skin.Predorsal scales with similar size as ventral and prepelvic scales (all cycloid).Cheek fully scaled (cycloid scales), with 6 -8 (n = 13) scale  First dorsal-fin spine about 1/3 to 1/4 of length of last dorsal-fin spine; spines increasing in length from first to last.Soft parts of dorsal and anal fin pointed, reaching to base of caudal fin.Caudal fin rounded.Pectoral fin rounded, originating on vertical line from first dorsal-fin spine, reaching back to about halfway to anal fin origin.Pelvic fin inserted posteriorly to pectoral axilla (at vertical line from third dorsal-fin spine), with acuminate tip, second ray longest, reaching about halfway to anal fin origin.
All teeth unicuspid, erect or slightly recurved.Teeth hyaline, red at tip in largest.Outer row teeth distinctly larger than inner teeth, and larger anteriorly than posteriorly; upper jaw with five inner rows, lower jaw anteriorly with three inner rows.
Microbranchiospines very small, easily overlooked, but present externally on first to fourth arches.Dentigerous plates present only on 4 th arches.
Ceratobranchial 5 triangular, strongly compressed dorsoventrally, slightly wider than long and with wide dentigerous area.Teeth short laterally, gradually increasing in size caudally and medially.Teeth in posterior row longest.Teeth round in cross section.Posterior margin row with 18 (n = 3), and in midline row with 7 (n = 3).
Coloration in alcohol (Fig. 1).No significant differences visible between males and females.Body light brown, darker dorsally and lighter ventrally, with 10 -12 trunk bars, above longitudinal stripe, from pectoral fin base to end of caudal peduncle.An interrupted dark brown longitudinal stripe from the end of caudal peduncle, extending along mid-portion of body to dorsal portion of opercle, crossing eyes, and reaching to tip of snout.Each trunk bar forming a blotch at junction with longitudinal stripe.Side of body with numerous dark brown spots (of a half body scale size), forming irregular, wavy, roughly horizontal lines, distributed below longitudinal stripe to ventral profile, and extending from posterior insertion of pectoral fin to end of caudal peduncle.No humeral ocellus.
Side of head with same coloration as trunk.Head without dots.Three dark brown head bars visible dorsally between posterior margin of orbits and posterior margin of operculum.Tips of upper and lower jaws darker than head.A narrow black or dark brown suborbital stripe, reaching about 4/5 of distance to preopercle with discontinuous distal portion (about 1/3).
Dorsal fin light brown, with irregular dark brown spots on entire fin.Anal fin with same coloration as dorsal fin.Caudal fin light brown, with irregular dark brown spots, more concentrated on proximal half.Posterior margin of caudal fin lighter than rest of fin.Ocellar spot on caudal fin base small, dark brown, with a pale brown or white ring.Pelvic and pectoral fins light brown.Etymology.The species name lucenai honours the ichthyologist carlos lucena for his contributions to the taxonomy and systematics of the genus Crenicichla.

Discussion
The most recent phylogenetic analyses (based on molecular data) revealed that Crenicichla is paraphyletic with respect to Teleocichla Kullander, 1988(Kullander et al., 2010;PiáleK et al., 2012).Thus, distinct monophyletic lineages within Crenicichla may be recognized as different genera (Kullander et al., 2010).The type species of the genus (C.macrophthalma hecKel, 1840) was recovered at a more basal position in the phylogenetic trees (Kullander et al., 2010;PiáleK et al., 2012), and is not particularly closely related to the species examined and discussed here.Nevertheless, in accordance with the current use (e.g.lucena & Kullander, 1992;Kullander & lucena, 2006;Montaña et al., 2008;casciotta et al., 2010;Varella et al., 2012), we include the new species C. lucenai sp.n. and its relatives in the genus Crenicichla, until the generic classification of crenicichlines is revised.
The melanin elements within the colour pattern are an important source of character states for the taxonomical delimitation of Neotropical cichlids (Kullander & Silfvergrip, 1991;Říčan et al., 2005;ottoni & cheFFe, 2009;ottoni & costa, 2009;ottoni, 2011;ottoni, 2013), and are often used as diagnostic traits to sepa-   rate closely related Crenicichla species (e.g.& lucena & Kullander, 1992;Kullander & lucena, 2006;2013;Montaña et al., 2008;Ploeg, 1991;Varella et al., 2012).The new species herein described is a member of the species group proposed by Kullander & lucena (2006) including C. maculata and C. punctata for having 59 -70 scales on E1 row, the presence of a row of blotches along the side of its body, and general colour pattern characters discussed in the diagnosis.The new species, C. lucenai sp.n., is clearly distinguishable from both C. maculata and C. punctata by its conspicuous pattern of dark brown irregular lines on body flanks (see diagnosis above).Such a pattern is unique within the genus.Hence, it may be treated as an apomorphic characteristic of the new species described here.
The area where these three species occur has undergone a complex tectonic history.It is associated with one of the six megadomes situated at the coastal area of eastern South America (riBeiro, 2006).Megadome uplifts and the erosive retreat of its margins had a significant influence in the drainages pattern and the development of the river basins (riBeiro, 2006).At present, C. luce nai sp.n. and C. maculata inhabit isolated hydrographic basins, separated by a mountain ridge.It is possible, however, that the upper Rio das Antas basin was once connected with the coastal rivers of the region where C. maculata is distributed.Therefore, it is not unlikely that C. lucenai sp.n. is more closely related to C. macu lata than to C. punctata.In overall appearance, C. luce nai sp.n. (Fig. 1) looks more similar to C. maculata (Fig. 3) than to its closest geographic neighbour C. punctata (Figs. 4 and 5).The dark spots present on the sides of the head in C. punctata (Figs. 4 and 5) distinguish this species from both C. maculata (Fig. 3) and C. lucenai (Fig. 1).
C. reticulata group by having higher number of depressible teeth rows, 3 -5 (versus often with fewer and more firmly implanted teeth); from C. macrophthalma hecKel, 1840 by having predorsal scales cycloid (versus predorsal scales ctenoid), normal size eyes (versus extremely large eyes) and lateral line scales with same coloration as adjacent flank scales (versus lateral line scales differently colored than adjacent flank scales); from the C. missio neira group by having preopercle margin serrated (versus smooth); from the C. scottii group by having maxilla just reaching to the vertical anterior of orbit (versus beyond the same vertical); from C. vittata by having 47 -76 E1 row scales (versus78 -85); from the species C. mucuryna and from the C. mandelburgeri species group by having wide bars on flank (versus narrow bars), and presence of spots on flank (versus absence); from C. iguapina, C. tin gui and C. lacustris by having a row of blotches on flank (versus a continuous lateral band).Crenicichla lucenai is distinguished from both C. punctata and C. maculata by having a conspicuous pattern of dark brown irregular lines extending from longitudinal stripe to ventral profile (versus irregular lines absent).In addition, C. lucenai is distinguished from C. puncatata by the absence of dots on sides of head (versus presence).
. Preopercle and interopercle naked.Opercle and subopercle fully scaled with diffuse pattern (all scales cycloids).Three scales between lateral lines.Upper lateral line not overlapping lower lateral line.Anterior upper lateral line scales larger than adjacent flank scales, remaining lateral line scales of nearly the same size as adjacent flank scales.Anterior lower lateral line scales slightly larger than adjacent flank scales, remaining lateral line scales of nearly the same size as adjacent flank scales.Dorsal, anal, pectoral, and pelvic fins without scales.Caudal fin squamation extending to nearly the middle of fin.

Figure 2 .
Figure 2. Distribution map of: Crenicichla lucenai sp.n., and type localities of the other species of the C. punctata group.
Description.Based on specimens larger than 103.0 mm SL.Measurements are given in table 1, counts in table 2; see Fig. 1 for habitus.Body elongate (mean of body depth 20.3 % SL).Caudal peduncle longer than deep.Snout moderately long, rounded in dorsal and ventral view, and moderately pointed in lateral view.Lower jaw prognathous.Maxilla reaching to vertical from anterior margin of orbit.Upper and lower lips of approximately the same width.Postlabial skin fold margin truncate.Orbit supralateral, not visible from below, chiefly in an-