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Description of a new species of annual fish, Maratecoara gesmonei (Cyprinodontiformes: Rivulidae) from the rio Xingu system, Amazon basin, Brazil.

Abstract

Maratecoara gesmonei n. sp., found on a temporary pool at a fluvial island in the middle rio Xingu, Para State, Brazil, is described herein. This is the first occurrence of this genus in the Rio Xingu drainage, Amazon basin. The new species differs from all congeners by its unique color pattern which lacks horizontal rows of small dark orange spots on the antero-dorsal portion of the flanks (vs. 2-3 on M. lacortei, 3 on M formosa and M splendida), or orange oblique bars on anteroventral portion of trunk (vs. 4-5 orange oblique bars on anteroventral portion of trunk in M formosa, 3-4 in M. splendida or a broad blotch in M. la-cone:). In addition, the new species can be diagnosed from congeners by its lower body depth (23.7-25.9% SL vs. 30.4-40.0% SL), lower caudal peduncle depth (13.015.5% SL vs. 17.1-21.6% SL), and lower number of vertebrae, 25-26 (vs. 27-28 in M. lacortei, 26-27 in M for-mosa, and 27 in M splendida).

Resumo

Maratecoara gesmonei n. sp., encontrada em uma poca tem-poraria em uma ilha fluvial no medio rio Xingu, estado do Para, Brasil, e aqui descrita. Esta 6 a primeira ocorrencia do genero na drenagem do Rio Xingu, bacia Arnazonica. A nova especie difere de todas as congeneres pelo padrao tinico de cor com ausencia de linhas horizontais e pequenas manchas laranjas escuras na porcao antero-dorsal dos flancos (vs. 2-3 na M lacortei, 3 em M formosa e M splendida), ou barras obliquas laranjas (vs. 4-5 barras obliquas laranja em M for-mosa, 3-4 em M splendida ou uma grande mancha em M lacortei). Adicionalmente, a nova especie pode ser diagnosti-cada de suas congeneres pela menor altura do corpo (23.725.9% SL vs. 30.4-40.0% SL), menor altura pedtinculo caudal (13.0-15.5% SL vs. 17.1-21.6% SL) e pelo menor mimero de vertebras, 25-26 (vs. 27-28 em M lacortei, 2627 em M formosa e 27 em M splendida).

Zusammenfassung

Beschrieben wird hier Maratecoara gesmonei n. sp, deren Vertreter in einem zeitlich begrenzten Tumpel auf einer Flussinsel im mittleren Rio Xingu im Bundesstaat Para in Brasilien entdeckt wurden. Es handelt sich urn den ersten Nachweis dieser Gattung im Einzugsgebiet des Xingu im Amazonasbecken, Brasilien. Die neue Art unterscheidet sich von alien anderen Angehorigen der Gattung durch das un-verkennbare Farbmuster: die waagerechten Reihen ldeiner, dunkel orangefarbener Flecken im vorderen Riickenbereich der Flanken fehlen hier (im Gegensatz dazu sind bei M la-cortei 2-3 Reihen, bei M. formosa und M. splendida 3 Fleck-enreihen vorhanden); auch fehlen hier die orangefarbigen schragen Streifen im vorderen bauchseitigen Teil des Rumpfes (bei MI formosa sind 4-5 schrage orangefarbene Streifen vorhanden, bei M splendida 3-4, bei M. lacortei emn breiter Fleck). Weitere Unterscheidungsmerkmale von den anderen Art der Gattung sind: eine geringere Korpertiefe (23,7-25,9 % der Standardlange SL im Gegensatz zu 30,440,0% der SL bei den anderen Arten), eine geringere Schwanzstieltiefe (13,0-15,5% der SL vs. 17,1-21,6% der SL) sowie eine kleinere Wirbelzahl (25-26 im Vergleich zu 27-28 bei M lacortei, 26-27 bei M formosa und 27 bei M splendida).

Resume

Maratecoara gesmonei n. sp., trouve dans une mare tern-poraire sur une Ile fluviale du moyen Rio Xingu, etat de Para, Bresil, est decrit ci-apres. C'est la premiere occurrence de cc genre dans le systeme du Rio Xingu, bassin de l'Amazone. La nouvelle espece se distingue de tous ses con-generes par un patron de coloration unique qui ne corn-prend pas de rangees horizontales de petites taches orange fonce sur la partie anterodorsale des flancs (contre 2-3 pour M lacortei, 3 pour M. formosa et M. splendida)ou des barres obliques orange sur la partie anteroventrale du tronc (contre 4-5 barres obliques orange sur la partie anteroven-trale du tronc pour M formosa, 3-4 pour M splendida ou une large tache pour M lacortei). En outre, la nouvelle espece peut etre separee de ses congeneres par sa plus faible hauteur du corps (23,7-25,9 % de LS vs. 30,4-40,0 de LS), une plus faible hauteur du pedoncule caudal (13,0-15,5 % de LS vs. 17,1-21,6 % de LS) et un nombre inferieur de vertebres, 25-26 (vs. 27-28 pour M lacortei 26-27 pour M formosa et 27 pour M splendida).

Sommario

Maratecoara gesmonei n. sp., trovata in una pozza temporanea su un'isola fluviale del medio corso del Rio Xingu, Stato di Para, Brasile, viene qui descritta. Questa e la prima segnalazione di questo genere nel sistema del Rio Xingu, bacino idrografico amazzonico. La nuova specie si differen-zia da tutte le congeneri per la particolare colorazione caratterizzata dall'assenza di righe orizzontali di piccole macchie di colore arancione scuro sulla porzione antero-dorsale dei fianchi (vs. 2-3 a M. lacortei, 3 su M formosa e M splendida) o barre oblique arancione sulla parte an-teroventrale del tronco (vs. 4-5 barre oblique arancione su parte anteroventrale del tronco in M. formosa e 3-4 in M. splendida o un'estesa macchia in M. lacortez). Inoltre, la nuove specie puo essere diagnosticata dalle congeneri per una minore altezza del corpo (23.7-25.9% SL vs. 30.440.0% SL), una minore altezza del peduncolo caudale (13.0-15.5% SL vs. il 17.1-21.6% SL) e un minor numero di vertebre, 25-26 (vs. 27-28 in M. Lacortei, 26-27 in M. formosa e 27 in M splendida).

INTRODUCTION

In the middle of the 1980's, annual fish enthusiasts collected three undescribed species of annual fish from the family Rivulidae in a temporary pool near the city of Aruana, located on the right bank of the rio Araguaia, Goias state, Brazil. These three new species were then temporarily identified by the codes GO-1, GO-2, and GO-3. All three species were very beautiful and distinct from what was then known to ichthyologists and aquarists at the time. One of them, GO-3, called a great deal of attention because it possessed a metallic blue coloration and long extensions on the dorsal and anal fins, recalling another species of annual fish, Terranatos dolichopterus Weitzman & Wourms, 1967, from the de Orinoco basin in Venezuela. Not until 1991 were these three species finally described as belonging to the genus Cynolebias Steindachner, 1876. The species that was initially designated with the code GO-3 was described as Cynolebias lacortei (Lazara, 1991).

Costa (1995a) erected a new genus, Maratecoara, and the species Cynolebias lacortei became the type species of this new genus. In the same work, a second species of Maratecoara was described, Maratecoara formosa Costa & Brazil, 1995. It originates from the city of Brejinho do Nazar, Tocantins state, and was found in an annual pool in the left bank of the rio Tocantins. More recently, Costa (2007), described one additional new species, Maratecoara splendida Costa, 2007, originating from an annual pool located close to the rio Canabrava, a tributary of the left bank of the rio Tocantins.

The genus Maratecoara belongs to the subfamily Rivulinae Myers 1925, tribe Rachoviini Costa, 1990 and the subtribe Plesiolebiina Costa, 1990. Within the subtribe Plesiolebiina, are also included the genera Plesiolebias Costa 1989, Pituna Costa 1989, Papiliolebias Costa 1998, and Stenolebias Costa 1995a. So far, the genus Maratecoara species is known exclusively from the rio Tocantins/ Araguaia basin in central Brazil. The new species described herein is the first found outside this system, at the middle rio Xingu basin, a the right bank tributary of the Amazon River. It is also the first species described from the Amazon forest domain - the other species are found in the savannah-like Cerrado domain.

MATERIAL AND METHODS

Measurements were taken point-to-point under the stereomicroscope with the digital caliper to the nearest 0.01mm, on the left side of the specimen whenever possible, following Costa (19956, 2007). Measurements are expressed as percentages of standard length (SL), except subunits of the head, which are recorded as percentages of head length (HL).

In the description, counts of vertebrae and pleural ribs were taken from cleared and stained (c&s) specimens, one male and one female paratypes, prepared according to Taylor & Van Dyke (1985). Terminology for frontal squamation follows Hoedeman (1958) and Costa (2006). For vertebral counts the caudal compounded centrum was counted as a single element. Osteological features included in the description are those considered phylogenetically informative by recent studies on Plesiolebiasiina (Costa 2007). Institutional abbreviations are LBP (LaboratOrio de Biologia e Genetica de Peixes, Universidade Estadual Paulista, Botucatu, Brazil), UNITAU (Universidade of Taubate, Taubate, Brazil), and ZUEC (Museu de Zoologia da Universidade Estadual de Campinas, Campinas, Brazil). Comparisons with congeners were based primarily on the literature (Costa, 2007).

Maratecoara gesmonei, n. sp.

(Figs 1-2; Table I)

Table I. Morphometric and meristic data for the holotype (H)
and paratypes of Maratecoara gesmonei. SD = Standard
deviation.

                       H  Para types    Females        MeantSD
                    Male    Male n=5        n=7

Standard length     23.4   18.5-20.4  17.9-24.5  19.4 [+ or -]
(mm)                                                      1.53

Percents of SL

Body depth          25.9   23.7-25.9  23.9-25.9  24.9 [+ or -]
                                                          0.87

Caudal peduncle     15.5   13.0-15.5  12.7-12.9  13.8 [+ or -]
depth                                                     1.26

Pre-dorsal length   66.8   63.2-67.8  60.1-66.8  65.9 [+ or -]
                                                          1.82

Pre-pelvic length   51.8   49.5-51.8  52.8-53.2   52. [+ or -]
                                                          1.12

Length of           14.5   13.5-14.5  11.3-13.3  13.2 [+ or -]
dorsal-fin base                                           l.17

Length of anal-fin  19.7   19.7-21.5  15.9-18.1  18.4 [+ or -]
base                                                      2.05

Caudal-fin length   41.4   34.1-41.4  31.9-37.2  36.0 [+ or -]
                                                          3.17

Pectoral-fin        23.3   23.3-27.2  19.1-23.8  23.7 [+ or -]
length                                                    2.27

Pelvic-fin length   14.8   12.2-14.8  12.7-14.9  13.5 [+ or -]
                                                          l.02

Head length         37.3   34.6-37.7  33.6-36.2  35.5 [+ or -]
                                                          1.47

Percents of HL

Head depth          62.5   62.5-64.6  63.0-64.7  06.7 [+ or -]
                                                          0.83

Head width          34.7   33.8-35.7  35.3-39.7  63.4 [+ or -]
                                                          2.37

Lower jaw length    13.8   13.8-15.4  12.8-14.4  14.3 [+ or -]
                                                          0.67

Eye diameter        31.9   30.8-34.3  30.8-33.3  32.0 [+ or -]
                                                          l.10

Counts

Dorsal fin            11          11         10

Caudal fin            24          25         24

Anal fin              14       14-15         14

Pelvic fin             7           7          7

Pectoral fin          13          13         13

Meristic

Scales in                      25-26         25
longitudinal
series

Scales in                          8          8
transversal series

Horizontal scales                 16         16
around caudal
peduncle


Holotype: ZUEC 7851, male 23.4 mm SL: Brazil, Para state, Sao Felix do Xingu, temporary pool at island in the middle of the rio Xingu 06[degrees]39'33.3" S 52[degrees]00'21.9" W; 17 May 2013, Ricardo Britzke & Mayler Martins.

Paratypes: LBP 18387, four males, 18.5-20.4 mm SL, six females, 17.9-24.5 mm SL, collected with the holotype. ZUEC 7852 (2 c&s), one male 19.6 mm SL and one female 18.8 mm SL, collected with the holotype.

Diagnosis: Males of Maratecoara gesmonei differs from the remaining congeners by a lower body depth (23.7-25.9% SL vs. 30.4-40.0% SL); lower caudal peduncle depth (13.0-15.5% SL vs. 17.121.6% SL); lower length of dorsal-fin base (13.514.5% SL vs. 14.9-19.6% SL); lower length of anal-fin base (19.7-21.5% SL vs. 20.3-27.2% SL); lower caudal-fin length (34.1-41.4% SL vs. 49.262.5% SL); lower pelvic-fin length (12.2-14.8% SL vs. 15.0-20.0% SL); lower head depth (62.564.6% HL vs. 92.2-112.4% HL); lower head width (33.8-35.7% HL vs. 54.9-65.8% HL); pectoral-fin posterior margin reaching vertical between base of the 1st and 2nd anal-fin rays (vs. 5thand 7th in M lacortei, 4th and 5th in M formosa, and 4th and 6th in M splendida); contact organs absent (vs. 3-5 on posterior margin of each scale on ventral portion of flanks in males in M lacortei, M formosa, and M splendida); anal-fin origin between pleural ribs of 12th and 13th vertebrae (vs.10th and 11th in M lacortei and M formosa, 10th and 12th in M splendida); lower number of caudal-fin rays, 24-25 (vs. 27-30 in M lacortei, 25-26 M formosa, and 25-27 M. splendida); lower number of pelvic-fin rays, 7 (vs. 8 in M lacortei, M formosa, and M splendida); lower number of transverse series of scales, 8 (vs. 9-10 in M lacortei, 9 in M. formosa, and M splendida); lower number vertebrae, 25-26 (vs. 27-28 in M lacortei, 26-27 M formosa, and 27 in M splendida). Additionally, males of Marate-coara gesmonei differs from males of the remaining species of the genus Maratecoara by presenting a body color pattern without horizontal rows of small dark orange spots on the antero-dorsal portion of flanks (vs. 2-3 in M lacortei, 3 in M for-mosa and M splendida), by lacking orange oblique bars on anteroventral portion of trunk (vs. 4-5 orange oblique bars on anteroventral portion of trunk in M formosa, and 3-4 in M splendida, and a broad blotch in M lacortez), and by presenting overall body coloration with irregular stains of orange and metallic blue (vs. overall body coloration metallic blue with horizontal orange spots in M la-cortei, metallic blue with horizontal and oblique orange spots in M formosa and M splendida).

Females of Maratecoara gesmonei differ from females of remaining species of the genus Marate-coara by presenting a lower body depth (23.925.9% SL vs. 28.2-34.3% SL); lower caudal peduncle depth (12.7-12.9% SL vs. 15.1-18.4% SL); lower length of dorsal-fin base (11.3-13.3% SL vs. 13.6-16.5% SL); lower length of anal-fin base (15.9-18.1% SL vs. 19.3-22.2% SL); lower caudalfin length (31.9-37.2% SL vs. 39.6-43.2% SL); lower pectoral-fin length (19.1-23.8% SL vs.24.5-29.7% SL); lower head depth (63.0-64.7% HL vs. 78.1-90.1% HL); lower head width (35.3-39.7% HL vs. 53.3-61.5% HL); and lower eye diameter (30.8-33.0% HL vs. 34.4-37.9% HL).

Descnplion: Morphometric data presented in Table I. Largest male examined 23.4 mm SL, largest female examined 24.5 mm SL. Dorsal profile slightly concave between head and base of dorsal-fin, slightly convex at caudal peduncle. Ventral profile gently convex from lower jaw to end of anal-fin base and nearly straight from latter point to caudal peduncle. Body moderately slender, greatest body depth at level of pelvic-fin base. Jaws short, prognathous, snout slightly pointed. Dorsal fin elongated in males, pointed, rays 7-8 much more elongated than remaining rays, filamentous, tips reaching at least distal margin of caudal fin when adpressed to the body; anal fin pointed in males, pointed, rays 78 much more elongated than remaining rays, filamentous, tips reaching distal margin of caudal fin when adpressed to body; dorsal and anal fins just slightly pointed in females. Caudal-fin lanceolate in males, middle caudal-fin rays filamentous, caudal-fin rounded in females. Pectoral-fin elliptical, distal margin reaching vertical between base of 1st and 2'd anal-fin rays in males, between urogenital papilla and anal-fin origin in females. Pelvic-fin elliptical, without filaments; tip of each pelvic fin reaching between base of 31rd and 4th anal-fin rays in males, reaching to between base of 1st and 2nd anal-fin rays in females. Pelvic-fin bases in close proximity medially. Dorsal fin origin on vertical between base of 3rd and 5th anal-fin rays, and between neural spines of llth and 12th vertebrae in males and 13th and 14tth in females. Anal fin origin between pleural ribs of 12th and 13th vertebrae. Dorsal-fin rays 10-11; anal-fin rays 14-15; caudal-fin rays 24-25; pectoral-fin rays 13; pelvic-fin rays 7.

Scales cycloid: No scales on caudal, dorsal and anal-fins bases. Frontal squamation F-patterned; E-scales not overlapping medially; scales arranged in regular transverse pattern. Longitudinal series of scales 2526; transverse series of scales 8; scale rows around caudal-fin peduncle 16. Contact organs absent.

Cephalic neuromasts: supraorbital 6+3, parietal 3, anterior rostral 1, posterior rostral 1, infraorbital 1 + 25, preorbital 4, otic 1, post-otic 2, supratemporal 1, median opercular 1, ventral opercular 1, preopercu-lar 14, mandibular 8, lateral mandibular 5. Two neuromasts on fin base. Total vertebrae 25-26.

Coloration in life (Figs 1-2). Males: Sides of body metallic blue and orange with irregular color distribution, without bars or lines, dorsally orange. Abdominal region clear. Sides of head metallic blue and orange. Jaws orange. Iris bright green; a black bar vertically crossing the eye. Dorsal-fin orange with metallic blue blotches. Anal-fin metallic blue with slightly orange stains. Caudal-fin orange with metallic blue spots. Pectoral-fins hyaline. Pelvic-fins metallic blue with an orange spot.

Females: Overall color light brownish gray, with longitudinal rows of pale brown dots on the sides of body. Abdominal region clear. Sides of head and jaws gray, with a pale greenish yellow hue. Iris yellow, with a dark gray vertical bar through center of eye. Fins hyaline.

Distribution: Currently only known from the type-locality, a temporary pool at an island from middle rio Xingu, Amazon basin, Para state, Brazil (Fig. 3).

Habitat (Fig. 4): The type-locality lies in a plateau area of Brazil (198 m a.s.1.), on an island in the middle of the rio Xingu, in front of the city of Sao Felix do Xingu. The island has a total area of about 465[m.sup.2] and a perimeter of about 3.160 km. It lies about 330 m from the left bank of the river and about 540 m from the right bank. The annual pool is very large with most of its area within a dense forest. Water temperature, at the depth of 25 cm was 26.5[degrees]C, whereas at the marginal area of the pool, at the depth of 5 cm, was 27.5[degrees]C. The average depth of the pool was 0.5 m, with the deepest portions about 1.20 m. Water color was dark and acidic, with pH 5.5. Other physic-chemical parameters were: electric conductibility 8 [micro]S, total hardness (GH) 0[degrees]dGH, carbonate hardness (KH) 0[degrees]dGH, dissolved iron (Fe) 0.5 mg/1, dissolved calcium (Ca) 0 mg/1, dissolved copper (Cu) 0 mg/1, dissolved phosphate (P[O.sub.4]) 0.1 mg/1, and ammonium/ammonia (N[H.sub.3]/N[H.sub.4]) 0 mg/l.

Other annual fish species collected syntopically were Pituna xinguensis and Plesiolebias altamira. The pool was beginning to dry up and the collectors also found juveniles of the following characiform fish species living syntopically: Moenkhausia xinguensis, M ceros, Jupiaba sp., Hyphessobrycon sp. Thayeria boehlkei; Serrassalmus sp., and Hoplias malabaricus. Other aquatic animals recorded at the site were tadpoles and freshwater crabs. Aquatic vegetation was not present and the bottom was composed of clay, leaves, sand and mud. Marate-coara gesmonei was always found in areas close the margin of the pool, nearby submerged trunks, at about 50 cm depth, while Pituna xinguensis was found in more shallow areas (about 5 cm deep) and Plesiolebias altamira in deeper areas (about 1 meter deep).

Etymology: The specific name gesmonei is treated as a patronym in gratitude and recognition to Ges-mone Fernandes Godoy, who discovered the species.

DISCUSSION

The genus Maratecoara is defined by the following synapomorphies: long dorsal and anal fins with tips extending beyond the posterior margin of caudal-fin; caudal-fin lanceolate, tip with two or three filamentous rays; long opercular membranes with blue iridescence, extending on to the anterior portion of pectoral-fin; and flanks metallic blue with orangish golden spots (Costa, 2007).

The discovery of annual pools in Sao Felix do Xingu increases the knowledge of distribution of annual fishes in the rio Xingu basin, which were so far only known to the neighborhoods of Altamira, where the Belo Monte dam is currently being build. The building of this dam will probably destroy the known habitat of the annual fishes that were so far known from the rio Xingu basin, i.e., Spectrolebias reticulatus, Plesiolebias altamira, Pitu-na xinguensis and Rivulus xinguensis. Fortunately, Pituna xinguensis and Plesiolebias altamira were found occurring syntopically with Maratecoara ges-monei, and consequently will not be extirpated by the building of the Belo Monte dam. Sao Felix do Xingu is about 384 km in a straight line from Al-tamira, and very likely, more habitats for annual fishes exist in the intervening area.

The type locality of Maratecoara gesmonei is considerably distant from the other known localities of Maratecoara species, i.e., 932 km from the type-locality of M lacortei, 815 km from the type-locality of M splendida and 640 km from the type-locality of M. formosa (Fig. 3). All Maratecoara species are only known from their type localities locations, with the exception for M lacortei, which is known from several sites at the rio Araguaia at the city of Aruand and its tributary, the rio das Mortes, and downstream into the Ilha do Bananal. Although the genus probably has a natural patchy distribution, this disjunctness may be a sampling artifact, both due to the relatively little fieldwork directed to the search of annual fishes, and also due to a possible great contraction of available habitat for the species of the genus and for annual fishes as a whole caused by the massive destruction of the Cerrado vegetation domain in Central Brazil, which is being replaced by large monoculture plantations.

The occurrence of the new species Maratecoara gesmonei in the rio Xingu basin might be associated to the neotectonic activity across the Transbrasil-iano Lineament area, a mega linear structure with about 2700 km, which begins in upper Paraguay River, crossing the Brazilian Shield to the northeast coast of Brazil, which also is hypothesized as the geological drive behind current ranges of other fish taxa occurring across shield-draining river basins of Central Brazil (Lima & Ribeiro 2011) (Fig. 5). These authors reported that the distribution of several fish taxa was probably shaped by several headwaters captures generated by neotectonic activity between the Pantanal depression and Araguaia-Tocantins depression, which probably allowed a interchange of fish populations among distinct river basins. Other genera of the subtribe Plesiolebiina, like Plesiolebias and Pituna, possess a similar distribution along the Transbrasilian Lineament and they also occur in the middle and lower rio Xingu region (Fig. 6).

ACKNOWLEDGEMENTS

We are grateful to Flavio C. T. Lima (ZUEC) and Claudio Oliveira (LBP) for curatorial assistance and Itamar Alves Martins from Universidade de Taubate (UNITAU) for laboratory support; Dr. Roger David Brousseau who reviewed the English version and Flavio C. T. Lima (ZUEC) for reading the manuscript and for offering useful suggestions.

REFERENCES

COSTA, W. J. E. M. 1995a. Two new genera and two new species of the neotropical annual fishes Plesiolebiasini (Cyprinodontiformes: Rivulidae), with studies on the relationships of the tribe. Revue Franfaise d'Aquariologie et Herpetologie 21: 65-74.

COSTA, W. J. E. M. 1995b. Pearl killifishes--the Cynolebi-atinae: systematics and biogeography of the neotropical annual fish subfamily. TFH, Neptune City, 128 pp.

COSTA, W. J. E. M. 2006. Descriptive morphology and phylogenetic relationship among species of the Neotropical annual killifish genera Nematolebias and Simp-sonichthys (Cyprinodontiformes: Aplocheiloidei: Rivuli-dae). Neotropical Ichthyology 4: 1-26.

COSTA, W. J. E. M. 2007. Taxonomy of the Plesiolebiasine killifish genera Pi tuna, Plesiolebias and Maratecoara (Teleostei: Cyprinodontiformes: Rivulidae), with descriptions of nine new species. Zootaxa 1410: 1-41.

LAZARA, K. J. 1991. Cynolebias lacortei, Cynolebias costai, and Cynolebias aruana: three new species of cloud fish from Brazil (Teleostei, Cyprinodontiformes, Rivulidae). Journal of the American Killifish Association 23: 139-152.

HOEDEMAN, J. J. 1958. The frontal scalation pattern in some groups of tooth carps. Bulletin ofAquatic Biology 1: 23-28.

LIMA, E C. T & A. RIBEIRO, C. 2011. Continental-scale tectonic controls of biogeography and ecology. Pp. 145164. In: Albert, J. S. & R. E. Reis (Eds.). Historical Biogeography of Neotropical Freshwater Fishes. Berkeley: University of California Press.

TAYLOR, W. R. & VAN DYKE, G. C. 1985. Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium 9: 107-119.

Dalton Tavares Bressane Nielsen (1), Mayler Martins (2) and Ricardo Britzke (3)

(1) Laboratorio de Zoologia, Departamento de Biologia, Universidade de Taubate, Pca Marcelino Monteiro 63, CEP: 12030-010, Taubate, SP, Brazil. E-mail: dnielsen@uol.com.br

(2) Instituto Federal Minas Gerais- Campus Bambui- Fazenda Varginha-Estrada Bambui-Medeiros Km 5, CEP: 38900-000, MG, Brazil. E-mail: maylermarins@yahoo.com.br

(3) Universidade Estadual Paulista, Instituto de Biociencias, Departamento de Morfologia, Rubido Jr. sin. CEP 18618-970. Botucatu, SP, Brazil. E-mail: britzke_r@yahoo.com.br

Received: 13 February 2014--Accepted: 03 April 2014
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Author:Nielsen, Dalton Tavares Bressane; Martins, Mayler; Britzke, Ricardo
Publication:aqua: International Journal of Ichthyology
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Geographic Code:3BRAZ
Date:Apr 1, 2014
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