Bryconamericus From Venezuela Paper

8/3/2019 Bryconamericus From Venezuela Paper

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15Animal Biodiversity and Conservation 31.1 (2008)

2008 Museu de Cincies NaturalsISSN: 1578665X

RomnValencia, C., Taphorn B., D. C. & RuizC., R. I., 2008. Two new Bryconamericus: B. cinarucoensen. sp. and B. singularis n. sp. (Characiformes, Characidae) from the Cinaruco River, Orinoco Basin, withkeys to all Venezuelan species. Animal Biodiversity and Conservation, 31.1: 1527.

AbstractTwo newBryconamericus: B. cinarucoense n. sp. andB. singularis n. sp. (Characiformes, Characidae) fromthe Cinaruco River, Orinoco Basin, with keys to all Venezuelan species.Herewe describe for the first timeBryconamericus cinarucoensen. sp. and Bryconamericus singularisn. sp., two new species of Characiformesfrom the Cinaruco River, Orinoco Basin in Venezuela. B. cinarucoensen. sp. is distinguished from all otherspecies of the genus in having: upper jaw extending beyond lower, maxilla short with only one or two teeth,cartilaginous rhinosphenoid extending to anterior part of prevomer, pelvic bone with cartilage along anterioredge, lateral line pores in straight line. B. singularisn. sp. is distinguished from congeners by having top ofhead flat, dentary with six or seven small unicuspid teeth, a dark lateral band extending from posterior edgeof humeral spot to midbase of caudal fin which widens behind dorsalfin origin, and in having fivesupraneurals which lack cartilage on the upper and lower extremities. Keys to aid identification of all knownVenezuela species are included. Bryconamericus motatanensis is placed in the synonymy of B. alpha.Previous reports of B. breviceps and B. heteresthes from Venezuela are misidentifications, and are hereconsidered as either B. cinarucoensen. sp., or another as yet undescribed species.

Key words:B. cinarucoensen. sp., B. singularisn. sp., Tropical fish, Taxonomy, Osteology, Teeth.

ResumenDos nuevosBryconamericus: B. cinarucoense sp. n. y B. singularis sp. n. (Characiformes, Characidae)del ro Cinaruco, cuenca del Orinoco, con claves para todas las especies de Venezuela. Se describendos especies nuevas de Bryconamericus de la cuenca del Orinoco en Venezuela: Bryconamericuscinarucoense sp. n. y B. singularis sp. n. (Characiformes, Characidae). B. cinarucoense sp. n. se

distingue de sus congneres por presentar la mandbula superior sobresaliente, la maxila corta y con unoo dos dientes, el rinoesfenoides cartilaginoso se extiende hacia la parte anterior del prevomer, el huesoplvico con cartlago a lo largo de su margen anterior, y por presentar los poros de la lnea lateral en lnearecta. B. singularis sp. n. se diferencia de las dems especies por presentar el extremo de la cabezaaplanado, seis o siete pequeos dientes unicspides en el dentario y una banda lateral oscura que seextiende desde el extremo posterior de la mancha humeral hasta la base media de la aleta caudal que seensancha en su parte posterior detrs del nivel del origen de la aleta dorsal, y por presentar cincosupraneurales sin cartlago en sus extremos superior e inferior. Se incluyen claves para la determinacinde las especies conocidas de Bryconamericus en Venezuela. Bryconamericus motatanensis se ubicacomo sinnimo de B. alpha. Citas previas de B. breviceps y B. heteresthes de Venezuela sonidentificaciones errneas, y aqu se considera como B. cinarucoense, u otra especie an no descrita.

Palabras claves: B. cinarucoensesp. n., B. singularissp. n., Pez tropical, Taxonoma, Osteologa, Dientes.

Two new Bryconamericus:B. cinarucoense n. sp. and B. singularis n. sp.(Characiformes, Characidae)

from the Cinaruco River, Orinoco Basin,with keys to all Venezuelan species

C. RomnValencia, D. C. Taphorn B. &R. I. RuizC.


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16 RomnValencia et al.

(Received: 18 VI 07; Conditional acceptance: 27 VIII 07; Final acceptance: 7 IX 07)

Csar RomnValencia & Raquel I. RuizC., Lab. de Ictiologa, Univ. del Quindo, A. A. 2639, Armenia,Quindo, Colombia.Donald C. Taphorn B, UNELLEZ, BioCentro, Coleccin de Peces, Museo de Zoologa,Guanare, Portuguesa, 3310, Venezuela.

Corresponding autor: C. RomnValencia. Email: [email protected]
mailto:%[email protected]:%[email protected]:%[email protected]:%[email protected]


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Animal Biodiversity and Conservation 31.1 (2008) 17

I n t roduc t ion

Species of the genus Bryconamericus Eigenmann(including Knodus Eigenmann) are predominantlysmall to mediumsized (usually 3050 mm maxi-

mum SL), silvery fishes with a humeral spot and adark lateral body stripe, silvery in life, that oftenextends onto the middle caudal rays (Gry, 1977;RomnValencia, 2002a, 2002b, 2003a, 2003b).Bryconamericus is complex genus of the familyCharacidae, with about fifty described species (Limaet al., 2003) widely distributed in a variety of fresh-water ecosystems in both the lowlands and high-lands of South and Middle America (Vari & Siebert,1990; Jimnez et al., 1998; RomnValencia, 2001,2002a, 2002b, 2003a, 2003b, 2003c, 2003d, 2005).Species of the genus are abundant in small brooksas well as along the banks of larger rivers, with highconcentrations of dissolved oxygen (ca. 8 mg/l) and

almost neutral pH (Jimnez et al., 1998; RomnValencia, 1998, 2000, 2002a; RomnValencia &Muoz, 2001). Bryconamericus species are knownfrom all major drainages in Venezuela (RomnValencia, 2003a) but have frequently beenmisidentified.

In recent studies by Lima & Zuanon (2004),Weitzman et al. (2005), Ferreira & Lima (2006), andFerreira & Carvajal (2007), the genus Knodus wasconsidered valid, although this was mainly for con-venience and based on the character of a scaledcaudal fin. However, there are no characters thatwould allow us to confidently separate Bryconamericusfrom Knodus; as pointed out by RomnValencia(2000, 2003a, 2005). In one key to Characidae,Planquette et al. (1996) attempt to differentiateBryconamericusand Knodus from Hemibryconby itsfewer than six teeth on the maxilla; they furthermorestate that the only difference between Bryconamericusand Knodus is the presence of scales on the caudalfin in the latter. There seems to be a consensus thatthe genus Bryconamericus, as currently defined, isnot monophyletic (Vari & Siebert, 1990; Malabarba &Malabarba, 1994; Silva & Malabarba, 1996; Malabarba& Weitzman, 2003; Silva, 2004). However, there isstill no published evidence that might indicate thatsome groups of Bryconamericus species are more

closely related to any other taxa (Vari & Sieber,1990; Malabarba & Kindel, 1995), nor proposals ofany new phylogenies. The difficulty in diagnosingBryconamericus and related genera was evidentwhen Malabarba & Malabarba (1994) describedHypobrycon maromba, but commented that it mightbe better to locate it in Bryconamericus. Serra &Langeani (2006) redescribed the type species ofBryconamericus (= B. exodon) and augmented thenumber of characters available for its diagnosis, butcommented that many of them may not representcharacters uniquely defining Bryconamericus.

Malabarba y Weitzman (2003) presented a hy-pothesis, placing Bryconamericusas a member of a

clade named "Clade A", supported by two synapo-morphies: four teeth in the innter premaxillary row,and ii,8 dorsalfin rays. Using molecular characters

Calcagnotto et al. (2005: fig. 6) found further supportfor the monophyly of Bryconamericus and indicatethat Knodus is its sister taxon, and that both areclosely related to Creagrutus and Hemibrycon. Alsousing molecular characters, RomnValencia &

VanegasRos (in press) recently proposed that thegenus Bryconamericus is a monophyletic group, atleast for the species from Central America. This wasalso proposed by Fink (1976). Furthermore, bothgroups (Fink, 1976; RomnValencia & VanegasRos, in press) present evidence that indicates thediversification of Bryconamericus in Central Americacan be explained by dispersion from northwesternSouth America. However, Lima et al. (2003) listsBryconamericusas insertae sedis in Characidae.

Taxonomically, Bryconamericus species fromCentral America have now been clearly resolved(RomnValencia, 2002a), but the South Americanspecies are still poorly understood; for most coun-

tries, for example Colombia (Magdalena drainage),Peru, Ecuador (Pacifico and Amazon drainages)and Bolivia, the available keys and species descrip-tions are of little use to determine the nominalspecies reported. We consider twelve species asvalid records from Venezuela in this report, includ-ing six from the Orinoco Basin. The description oftwo new species of Bryconamericus from theCinaruco River in Venezuela adds to the first au-thor's ongoing revision of the species in northernSouth America, and is further proof of the asyetundocumented biodiversity of the genus. We pro-vide keys to aid identification of all Venezuelanspecies for the following regions: Maracaibo Basin,Caribbean coastal drainages, Orinoco Basin, andthe Rio Negro drainage in Amazonas state.

Mater ia l and methods

Twentyone measurements were taken with digitalcalipers, recorded to hundredths of millimeters andexpressed in most cases as percentages of stand-ard or head length (table 1). Nine counts weremade using a stereoscope with a dissection needleto extend the fins. Counts and measurements weretaken from the left side of specimens when possi-

ble, basically following the guidelines in Vari &Siebert (1990).Observations of bones and cartilage were made

on cleared and stained specimens prepared ac-cording to techniques outlined in Taylor & Van Dyke(1985) and Song & Parenti (1995). Bone nomencla-ture follows Weitzman (1962), Vari (1995), RuizC.& RomnValencia (2006). Specimens are depos-ited in the Auburn University Museum Fish Collec-tion, Auburn, Alabama (AUM), the Museo deBiologa, Instituto de Zoologa Tropical, UniversidadCentral de Venezuela, Caracas (MBUCV), Museode Historia Natural La Salle, Caracas (MHNLS), theMuseo de Ciencias Naturales de la UNELLEZ

Guanare, Venezuela (MCNG), the IchthyologyLaboratory at the Universidad del Quindo, Armenia,Colombia (IUQ) and in the Instituto de


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Table 1. Morphometric and meristic data of Bryconamericus singularis n.sp. and B. cinarucoense. n. sp.(Standard and total length in mm, averages in parenthesis.)

Tabla 1. Datos morfomtricos y mersticos deBryconamericus singularis sp. n. yB. cinarucoense. sp. n.

(Longitud estndar y total en mm, medias entre parntesis)

B. singularis B. cinarucoense

Holotipo Paratipos Paratipos Holotipo

Standard length 33.41 27.3133.57 (30.50) 19.7533.57 (26.81) 28.98

Total length 41.07 35.4041.55 (38.82) 23.6242.88 (32.96) 36.76

Percentages of SL

Body depth 19.47 19.4725.99 (23.58) 16.9725.94 (20.61) 19.43

Snoutdorsal fin distance 52.05 47.1952.29 (49.71) 40.9456.78 (50.41) 53.00

Snoutpectoral fin distance 26.58 21.9930.07 (26.33) 19.4429 .45 (24.76) 25.85

Snoutpelvic fin distance 49.75 42.2552.10 (47.84) 39.9453.56 (46.75) 46.69

Dorsalpectoral fin distance 36.55 33.7840.62 (37.09) 29.9140.97 (36.51) 32.15

Snoutanal fin distance 66.96 63.3368.98 (65.65) 48.4665.62 (59.77) 60.08

Dorsal finhypural distance 46.96 46.8556.23 (51.46) 35.0354.10 (47.85) 47.70

Dorsalanal fin distance 27.49 21.4837.31 (25.09) 16.5444.70 (22.23) 19.57

Dorsalfin length 21.88 18.6628.38 (25.59) 14.426.60 (20.65) 19.60

Pectoralfin length 19.84 17.7228.01 (22.85) 15.6326.0 (20.8) 23.50

Pelvicfin length 16.55 13.8522.56 (18.45) 10.6820.10 (15.24) 14.84

Analfin length 18.26 17.4520.18 (18.63) 13.1121.06 (16.49) 18.77

Caudal peduncle depth 8.8 7.8810.55 (9.28) 6.059.54 (8.22) 8.45

Caudal peduncle length 10.72 8.6416.80 (11.77) 8.322.81 (14.68) 14.53

Head length 21.59 21.5927.38 (24.06) 18.7730.85 (24.42) 23.78

Percentages of HL

Snouth length 21.63 21.6329.52 (25.35) 18.4934.55 (25.71) 26.27

Orbital diameter 53.09 39.5953.09 (46.79) 35.7654.74 (44.79) 45.28

Postorbital distance 29.17 29.1737.50 (33.15) 27.7249.64 (36.31) 30.19

Maxilla length 32.93 18.2732.93 (27.30) 21.3437.54 (29.75) 21.80

Interorbital distance 28.40 28.4038.05 (33.87) 25.5739.46 (32.63) 32.51

Mandible superior superior 19.87 19.8731.06 (25.60) 19.2030.70 (25.03) 20.14

Lateralline scales 31 3032 3537 36

Scale row between dorsalfin origin and lateral line

5 56 45 5

Scale rows between analfin origin and lateral line

4 4 34 4

Scale rows between pelvicfin and lateral line

4 4 34 4

Predorsal median scales 10 10 1011 10

Dorsalfin rays ii,8 iiiiv,8 iiii,78 iii,7

Analfin rays iii,17 iii,1718 iiiiv,1821 iii,20

Pelvicfin rays ii,8 ii,6 ii,6 ii,6

Pectoralfin rays ii,10 ii,1011 ii,1011 ii,11


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Investigaciones Biolgicas "Alexander VonHumboldt", Villa de Leyva, Boyac (IaVH).

In the lists of paratypes, the number of individu-als is given in parentheses immediately after thecatalog number, which is followed by the range ofStandard Length (SL) in mm for that lot; for exam-ple: MCNG 53000 (23 ex.) 19.3425.44, indicates23 individuals in lot MCNG 53000, with the small-est fish 19.34 mm SL and the largest 25.44 mm SL.All collections were made in Venezuela. If no meas-urements are presented, the paratypes were notmeasured.

Comparative material (all from Venezuela)

Bryconamericus alphaEigenmann, Henn and Wilson1914 (see RomnValencia, 2003a)Bryconamericus breviceps Eigenmann 1908 (seeRomnValencia, 2003d)Bryconamericus heteresthesEigenmann 1908(seeRomnValencia, 2003a)Bryconamericus orinocoense RomnValenciaRomnValencia 2003d

Holotype: MBUCV 29464, 28.1 mm SL; AmazonasState, Ro Orinoco 0.5 km upstream from Esmeraldas(aprox. 253 06'' N, 6458' 06'' W); 12 III 1987.

Paratypes: collected with holotype: IUQ 433(9 ex.) MBUCV 25834 (26); MBUCV 6055 (1 ex.)Amazonas, La Esmeralda, Cao Cadabaudi,23 XI 1969. MBUCV 19395 (3 ex.); Amazonas, RoMavac near base camp; MBUCV 21658 (3 ex.),Amazonas, Ro Cataniapo, 10 XI 1989.

Resul ts

Bryconamericus cinarucoensen. sp. (fig. 1, table 1)

Holotype: MCNG 52002, 28.98 mm SL: Venezuela,

Orinoco River basin, Cinaruco River, Apure State,Pedro Camejo County, sand beach, 6 32' 55'' N;67 24' 58'' O, 21 V 1999 A. Arrington, C. Garca.

Paratypes: all from Venezuela, Orinoco River ba-sin, Apure State, Cinaruco River: MCNG 34679(6 ex.), 17.4434.26 mm SL, beach in front of LagunaLarga, Apr1997, coll. David Jepsen, DouglasRodrguez; MCNG 34705 (31 ex.) 12.8020.08, beachnear Laguna Larga, 4 IV 97, coll. David Jepsen andDouglas Rodriguez; MCNG 39174 (2 ex.), 31.6633.62, beach above Laguna Larga, 11 VI 99, A. andJ. Arrington; MCNG 39248 (27 ex.) 30.1136.87,beach below Laguna Larga, 11 VI 99, coll. A. & J.Arrington; MCNG 41277 (88 ex.) 17.5335.63, beach,19 IV 1999, coll. A. & J. Arrington; MCNG 41292(26 ex.) 23.2732.64, beach, 19 VI 1999, coll. A. &J. Arrington; MCNG 41638 (247 ex.), 18.8637.26,IUQ 526 (16 ex.), IUQ 527 (3 ex.) (C & S), 25.228.4collected with holotype; MCNG 41677 (13 ex.) 18.0632.81, beach 7, 21 V 1999, coll. A. Arrington & C.Garca; MCNG 44438 (10 ex.) 30.9436.79, beachupstream from Laguna Larga, 20 VI 2001, coll. C.Layman; MCNG 44496 (27 ex.) 29.6735.61, beachupstream from Laguna Larga, C. Layman.

Nontype material examinedFrom Venezuela, Orinoco River basin, Apure State,Cinaruco River: MCNG 39175 (4 ex.), above LagunaLarga, 11 VI 1999, coll. A. & J. Arrington; MCNG39221 (1 ex.), in front of Laguna Larga, 11 VI 1999,

coll. A. & J. Arrington; MCNG 40549 (178 ex.),Laguna Espinar, 18 Mar 1999, coll. A. & J. Arrington;MCNG 40842 (64 ex.), Cinaruco River 24 III 1999,coll. A. & J. Arrington; MCNG 41147 (5 ex.), beach4, 15 IV 1999, coll. A. & J. Arrington; MCNG 41575(14 ex.), beach 2, 21 V 1999, coll. A. Arrington & C.Garca; MCNG 45013 (1 ex.), 17 V 1999, coll. A.Arrington & C. Garca; MCNG 45014 (1 ex.),downriver from Laguna Larga, 11 VI 1999, coll. A.& J. Arrington.

From Guyana Essequibo River basin: AUM 38844(54 ex.), Takuku River 3.77 km SSW LethemRupununi, latitude 03.35500, 1 XI 2003, coll. J. W.Armbruster et al.; AUM 39017 (34 ex.), Essequibo

river at Yukanopito Falis, 44.5 km SW mouth ofKuyuwini River, latitude 01.91461, 9 XI 2003, coll. J.W. Armbruster et al.; AUM 44686 (4 ex.), Pirara

Fig. 1. Bryconamericus cinarucoensen. sp.: holotype, MCNG 52002, 28.98 mm SL.

Fig. 1. Bryconamericus cinarucoense sp. n.: holotipo, MCNG 52002, 28,98 mm LE.

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River, at Pirara Ranch, latitude 03.62517, 26 XI 2005,coll. L. S. de Souza et al.; AUM 38952 (18 ex.),Araquai Creek 77.3 km SSE Lethem Rupununi,latitude 02.76261, coll. J. W. Armbruster et al.,15 XI 2003; AUM 44948 (30 ex.), Guyana, Ireng

River, 6.9 km WSW Karasabal, latitude 04.01957,1 XI 2002, coll. J. W. Armbruster; AUM 38104 (5 ex.),Creek and Kuyuwini River 28.0 km E KuyuwiniLanding Rupununi, latitude 02.04747, 5 XI 2003, coll.J. W. Armbruster et al.

DiagnosisBryconamericus cinarucoense n. sp. is distin-guished from congeners by having the upper jawextending beyond lower, maxilla short with one ortwo teeth, cartilaginous rhinosphenoid extendingto anterior part of prevomer, pelvic bone withcartilage along anterior edge, and lateral line poresin a straight line. It differs from B. subtilisform

RomnValencia (RomnValencia, 2003b), themost similar Venezuelan species, in having fewerpored lateral line scales (3537 vs. 3839), morescales below lateral line to origin of pelvicfin (45 vs. 23), more branched analfin rays (2021vs. 1718), and pelvicfin rays usually ii,6 vs. i,7in B. subtilisform [see also: table 1 (this paper),and table 1 (RomnValencia, 2003b), and keysincluded below].

DescriptionBody slender and elongate (mean maximum bodydepth about 20% SL). Area above orbits flat. Dorsalprofile of head and body oblique from thesupraoccipital to dorsal origin and from the lastdorsalfin ray to the base of the caudal fin. Ventralprofile of body convex from the snout to the base ofanal fin. Caudal peduncle laterally compressed.Head and snout short, mandibles not equal, theupper longer than the lower; mouth terminal, lipssoft and flexible and not covering the outer row ofpremaxillary teeth; ventral border of the upper man-dible straight; posterior edge of the maxilla reach-ing anterior edge of orbit; opening of posteriornostrils vertically ovoid; opening of anterior nostrilswith a membranous flap. Dorsal surface ofmesethmoid covered with cartilage, which extends

all along the sensorial canal.Four or five infraorbitals with laterosensorial ca-nal present; first infraorbital thin and narrow, extend-ing between the dorsal edge of maxila and lateralethmoid, second infraorbital short and wide, notcompletely covering the dorsal part of the anguloarticular, anterior part squared off and exposed tosurface. Third infraorbital the widest and longest, itsventral border in contact with the preopercle; fourthand fifth infraorbitals short and narrow, covering thehyomandibular. Supraorbital present. Premaxilla withascending lateral process and two rows of teeth;external row with six tricuspid teeth arranged in astraight line except for the first proximal tooth which

is a little out of line; internal row with four teeth, eachwith three to five cusps, the central cusp largest.Maxilla short, the posterior edge not reaching ante-

rior edge of the second or third infraorbital. Maxillawith one or two teeth with three or four cusps each.Dentary with four large pentacuspid teeth with thecentral cusp largest, followed by six or seven smallteeth, the first tricuspid and the last two unicuspid.

Rhinosphenoid osseous, with cartilaginous borderand attached to orbitosphenoid by cartilage andextending to anterior edge of prevomer. Orbitosphe-noid wide, short and united to pterosphenoid by aband of cartilage. Palatine united with parasphenoidby cartilage.

Dorsal fin with oblique dorsal edge, the secondray simple and the first two branched rays thelongest. Radial and proximal pterygiophores of allrays of the dorsal fin inserted between the neuralspines 1118. Four to six supraneurals presentbetween the head and the anterior part or thedorsal fin, with cartilage on the upper and loweredges. Pterygiophores of the anal fin completely

cartilaginous, with just a small ossification of thethree proximal anterior pterygiophores.

Pectoral girdle with a pointed dorsal process abovethe cleithrum that surpasses the entire supracleithrum,which is joined to the postemporal. Cartilage presentat the union of scapular with the internal surface ofthe supracleithrum. Four proximal radials. Pelvicbone short, straight, blunt and with cartilage at theanterior tip, its posterior projection extending betweenthe junction of the two rows of pelvic rays.

Pelvic fin long, but not reaching the origin of theanal fin. Caudal fin not scaled, forked with shortpointed lobes, 910/910 principal caudal rays.Cartilage present at the basal part of the lastcaudal vertebra and the urostyle. Lateral line with3536 pored scales that extend in a straight linefrom the supracleithrum to the hypurals. Total ver-tebra 35. No sexual dimorphism was observed.

DistributionThis species is known from the Cinaruco River ofsouthern Apure State (fig. 3) and from the EssequiboRiver Basin of Guyana, and probably extends intosimilar rivers throughout the Orinoco Basin in Ven-ezuela and Colombia.

Etymology

Bryconamericus cinarucoense n. sp. is named forthe Cinaruco River of southern Apure Sate, wherethe type series was collected.

HabitatBryconamericus cinarucoense n. sp. was collectedalong shore over sandy substrates in the main-stream of rivers, as well as tributaries with flow. Thetransparency of the teacolored water is usuallymoderate to high, total dissolved solids and conduc-tivity are very low, and pH is usually slightly acidic.

Bryconamericus singularis n. sp. (fig. 2, table 1)

Holotype: MCNG 54500, 33.41 mm SL, Venezuela,Apure state, Orinoco River basin, Cinaruco River,


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beach (6 32.33 N 67 25.35 W), 20 II 1999, coll.D. A. Arrington.

Paratypes: All from Venezuela, Apure State,Cinaruco River. Taken with holotype: IUQ 541 (3 ex.cleared and stained), MCNG 39659 (137 ex.); MCNG39177 (14 ex.) 28.1932.41, beach upstream fromLaguna Larga (6 33.15 N 67 25.45 W),11 I 1999; MCNG 39194 (6 ex.) 25.7633.60, beachupstream from Laguna Larga (6 32' 20'' N 6724' 81'' W), 11 VI 1999; MCNG 39232 (3 ex.)29.4430.69, beach downstream from Laguna Larga(6 32.52 N 67 24.52 W), 11 VI 1999; MCNG

39245 (8 ex.) 27.7332.78, beach downstream fromLaguna Larga (6 32.50 N 67 24.08 W),11 I 1999; MCNG 39440 (10 ex.) 25.2631.13, beachdownstream from Laguna Larga (6 33.37 N 67 2.55 W), 6 VI 1999; MCNG 40060 (28 ex.)20.3023.36, & MBUCV 33029, 22.2325.45, RoCinaruco, Laguna Espier, Dtto. Pedro Camejo, Edo.Apure, 17 II 1999; MCNG 40149 (15 ex.) 23.9334.03, beach (636.32 N 6714.87 W), 1 II 1999;MCNG 40374 (12 ex.) 26.2430.83, LagunaEstrechura (632.25 N 6716.95 W), 16 III 1999;MCNG 40932 (2 ex.) 29.3929.83, Laguna Espier,12 VI 1999; MCNG 41014, (6 ex.) 26.6232.01,Laguna Estrechura, 13 VI 1999; MCNG 41640

(18 ex.) 29.0237.85, beach 5 (6 32.92 N 6724.97 W), 21 VI 1999; MCNG 41657 (17 ex.)29.4438.62, beach 6 (632.52 N 6724.52 W),21 V1999.

Nontype material examinedAll from Venezuela, Apure State, Cinaruco River:MCNG 39149 (1 ex.) beach upstream from LagunaLarga (6 38.05 N 67 26.05 W), 11 VI 1999;MCNG 39562 (2 ex.) beach, (6 32.92 N 724.97 W) 2 X 1999; MCNG 39565 (4 ex.) beach(6 32.50 N 67 24.97 W), 10 II 1999; MCNG39686 (5 ex.) beach (633.05 N 6726.62 W),2 II 1999; MCNG 39721 (36 ex.) beach (632.92 N

6724.97 W), 20 II 1999; MCNG 39770 (19 ex.)beach (6 32.52 N 67 24.52 W), 18 II 1999;MCNG 39801 (42 ex.) beach (6 32.50 N

6724.08 W), 18 II 1999; MCNG 39902 (17 ex.)beach (6 33.37 N 67 22.55 W), 16 II 1999;MCNG 39946 (4 ex.) Laguna Espier (632.80' N 6725.90 W), 16 II 1999; MCNG 40284 (7 ex.),beach (6 33.32 N 67 14.87 W), 15 III 1999;MCNG 40312 (16 ex.) Laguna Guayaba (634' 83''N 67 13' 84'' W), 16 III 1999; MCNG 40403

(35 ex.) Laguna Estrechura (6 32.32 N 6714.87 W), 16 III 1999; MCNG 40578 (1 ex.)beach (6 32.92 N 67 24.97 W), 18 III 1999;MCNG 40588 (1 ex.) beach (6 32.92 N 6724.97 W), 18 III 1999; MCNG 40769 (1 ex.)

beach (6 32.53 N 67 24.82 W), 24 III 1999;MCNG 40792 (88 ex.) beach (6 32.33 N 6725.35 W), 24 III 1999; MCNG 40809 (40 ex.)beach (6 32.33 N 62 25.35 W), 24 III 1999;MCNG 40829 (1 ex.) beach (6 32.33 N 62 25.35 W), 24 III 1999; MCNG 41455 (3 ex.)beach (6 32.50 N 67 24.08 W), 17 VI 1999;MCNG 41459 (1 ex.) beach (6 32.50 N 67 24.08 W), 17 V 1999; MCNG 41562 (1 ex.)beach (6 32.53 N 67 24.28 W), 21 VI 1999;MCNG 41572 (1 ex.) playa 2 (6 32.53 N 6725.45 W), 21 V 1999.

Diagnosis

Bryconamericussingularisn. sp. differs from con-geners in having the top of the head flat, thedentary with six or seven slight unicuspid teeth; adark lateral band extending from posterior edgeof the usually diffuse humeral spot to the base ofcaudal fin, and that widens and strengthens inintensity posteriorly at point beneath dorsal finorigin; five supraneural bones without cartilage oneither upper or lower edges and without cartilageon all structures.

DescriptionBody elongate, dorsal profile of head and anteriorbody rising from snout to dorsalfin origin, inclined

downwards from last dorsalfin ray base to base ofcaudal fin. Ventral profile of body straight fromsnout to base of anal fin. Head and snout short;

Fig. 2. Bryconamericus singularisn. sp.: holotype, MCNG 33.41 mm SL.

Fig. 2. Bryconamericus singularis sp. n.: holotipo: MCNG 33,41 mm LE.

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jaws equal in length; mouth terminal; lips soft andflexible, not covering the external row of premaxillaryteeth; premaxila with one ascendent processes thatarticulate with mesethmoid and a lateral process thatsupports the teeth and articulates laterally with theascendent process of the maxilla; posterior end ofmaxilla extends beyond anterior edge of orbit. Fiveinfraorbitals with sensory canal present; first infraorbitalthin, extending between the dorsal border of maxilla

and lateral ethmoid, the second long, covering thedorsal portion of anguloarticular and anterior part ofquadrate, third infraorbital wider, its posteroventralborder in contact with preopercle, fourth and fifthinfraorbitals short and narrow, covering thehyomandibular. Supraorbital absent. Premaxilla withtwo rows of teeth; the outer row with four tricuspidteeth with bases arranged in straight line. Internal rowwith five tri to pentacuspid teeth, with the centralcusp much longer than rest. Maxilla with one or twotricuspid teeth. Dentary with four large front teeth,those at middle pentacuspid, those on sides tricuspid,all with central cusp much the larger, followed by sixor seven small unicuspid teeth.

Along the ventral portion of the supraoccipitalprocess there are one or two foramens above thesupraoccipital canal that communicate with the neural

complex; dorsalmost portion of neural complexextending as two small apophyses that continueventrally as a canal. Rhinosphenoid osseous unitedto orbitosphenoid by a thin osseous plate. Osseousrhinosphenoid united to orbitosphenoid by thin laminarbone. First two branched dorsalfin rays longer thanrest. Proximal dorsal pterygiophores inserted betweenneural spines 910 and 1617. Five supraneuralspresent, lower and upper ends without cartilage.

Cartilage absent from the union between scapulawith the internal surface of the cleithrum, and ingeneral from all its structures. Pectoral girdlearticulated posterolaterally with cranium by fusionwith the supracleithrum and ventral end ofposttemporal bone; united to dorsal edge of cleithrum.Cleithrum located beneath ventral edge of opercle,three postcleithrals present above posterior edge ofpectoral girdle, first postcleithral posterior to union ofpostcleithrum and posttemporal, second and thirdpoiscleithrals united below with cleithrum whichextends over the pectoral rays. Three or four proxi-mal radials. Pelvic fin short, its tip not reaching analorigin. Pelvic bone long and straight, its lateral edge

convex, internal concave, and located parallel tocentral axis of body; ischial process with a short,straight pointed apophysis.

Fig. 3. Geographic distribution of B. cinarucoensen. sp. () and B. singularisn. sp. () in Venezuela.

Fig. 3. Distribucin geogrfica deB. cinarucoense sp. n. () y deB. singularis sp. n.() en Venezuela.

Caribbean Sea

Brazi l

Colombia

Guyana

Orinoco

Orinoco

Cuyun

Negro 100 0 100 200 300 400 km

12

10

8

6

4

2

0

12

10

8

6

4

2

0

7 4 7 2 7 0 6 8 6 6 6 4 6 2 6 0 5 8

7 4 7 2 7 0 6 8 6 6 6 4 6 2 6 0 5 8


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portion of anal fin base with concentration ofmelanophores, some specimens with melanophoresoutlining the muscle bundles, forming chevrons.Head dark and countershaded.

DistributionKnown from the Cinaruco River, Orinoco Basin,Apure State, Venezuela (fig. 3).

DietIt is omnivorous, feeding on aquatic insects, snails,seeds and aquatic plants. Stomach contents of fivefish included: Hemiptera: Vellidae: Microbelia (5.8%by Number, 33.3% by Frequency of Occurrenceand 5.41% Volume) insect parts (100% F, 32.4% V),Mollusca: Bivalvia (47% N, 100% F & 4.05% V),seeds (41.1% N, 100% F & 4.05% V), plant stems(5.8% N, 33.3% F & 1.35% V) and vegetable mat-ter (66.6% F & 32.4% V).

EtymologyThe name refers to the singular and striking aspectof this new species.

Caudal fin forked with long pointed lobes. Principalcaudal rays 10/10, no scales at base. 3032 poredscales in the lateral line, which ends on the caudal fin.Pores of the lateral line forming a gentle curve fromfirst to seventh scales, the rest in straight line. Total

vertebrae 33. No sexual dimorphism observed.

Color in alcoholBody light yellow, darker on dorsum. Lateral portionof body with a dark band behind humeral spot thatextends to the base of the caudal fin, and thatwidens at point beneath dorsal fin origin. Guaninepresent dorsally and laterally and on opercle inmany specimens. Humeral spot present but usuallydiffuse, about same height as pupil of eye and notusually extending dorsally beyond lateral stripe.Exposed edges of scales on dorsum and uppersides edged with black. Tips of caudal lobes dark,light yellow color of body extending onto central

caudal rays, forming light spot at base of caudal finthat is bordered above and below by black. Dorsal,anal, pectoral and pelvic fins hyaline. Anal fin lightlypigmented at tips of rays, body above anterior

Key to the species of Bryconamericus from the Cinaruco River.

Clave para las especies deBryconamericus del ro Cinaruco.

1 Sides of body with a welldefined, dark, lateral stripe;humeral spot absent or only weakly developed (diffuse) andnot usually vertically elongated; lateral scales 33 or fewer 2

Sides without wide, dark, lateral stripe; humeral spotstrongly developed, vertically elongate;lateral scales 35 or more Bryconamericus cinarucoense

2 Unbranched dorsal fin rays usually ii; unbranchedanalfin rays usually iv; length of maxilla 3239%of head length, mean 35%; body short and stocky,its greatest depth 2530% of standard length,mean 27% Bryconamericus orinocoense

Unbranched dorsal fin rays usually iiiiv; unbranchedanalfin rays usually iii; length of maxilla in head length

1832%, mean 27%; body long and slender,its greatest depth 1926% standard length, mean 24% Bryconamericus singularis

Key to the species of Bryconamericus from tributaries of the Gulf of Paria.

Clave para las especies de Bryconamericus de los afluentes del golfo de Paria.

1 Lateral scales 3536; total anal-fin rays 3031;maxilla with 34 teeth Bryconamericus lassorum

Lateral scales 3942; total anal-fin rays 2629;maxilla with 58 teeth Bryconamericus yokiae


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Key to the species of Bryconamericus from the Caribbean Coastal drainages.

Clave para las especies deBryconamericus de las vertientes de la costa caribea.

1 Total anal rays (simple plus branched) fewer than 21;small red or yellow dot present on upper caudal peduncle Bryconamericus cismontanus

(Caribbean drainages of Falcon,Lara, Yaracuy and other states)

Total anal rays more than 20 22 Lateral scales fewer than 35; teeth on maxilla multicuspid

with all cusps of equal length Bryconamericus charalae(endemic to Rio Aroa drainage, Lara)

Lateral scales more than 34; teeth on maxilla multicuspidwith central cusp longer than others 3

3 More than four small teeth behind major series of largeteeth on dentary; three to four unbranched analfin rays;body elongate (greatest body depth 26.6% SL) Bryconamericus loisae

(Zulia, Falcn, Lara, Carabobo,Yaracuy)

Fewer than three small teeth behind major series oflarge teeth on dentary; more than four unbranched analfinrays; body shorter and higher (greatest body depth 30.1% SL) Bryconamericus alpha

(Aragua, Miranda, Anzotegui)

Key to the species of Bryconamericus from the Lake Maracaibo drainage.

Clave para las especies deBryconamericus de las vertientes del lago Maracaibo.

1 Anal fin rays iii,13 to iii,16; scales from lateral lineto anal fin base 23 Bryconamericus meridae

Anal fin rays iiiv, 1830; scales from lateral lineto anal fin base 4 or more 2

2 Anal fin rays v, 1823; lateral scales 3238; body shorterand stocky, its greatest body depth 30.07% SL Bryconamericus alpha

Anal fin rays iii, 2430; lateral scales 3841; body longand slender, its greatest body depth 26.64% SL Bryconamericus cf loisae

Key to the species of Bryconamericus of the Apure and Arauca drainages.

Clave para las especies deBryconamericus de las vertientes de Apure y Arauca.

1 Sides without dark lateral stripe that continues onto centralcaudalfin rays; caudal fin usually with a vertically oriented,crescentshaped dark blotch at base Bryconamericus cinarucoense

Sides with a dark lateral stripe that continues onto centralcaudalfin rays 2

2 Five or more small teeth behind main series on dentary;body elongate (maximum body depth 26.6% SL);no small red or yellow dot on upper caudal peduncle Bryconamericus loisae

Four or fewer small teeth behind main series on dentary;body not as elongate (maximum body depth 273% SL),small red or yellow dot on caudal peduncle present or absent 3

3 Branched analfin rays 20 or more; no red or yellowdot on caudal peduncle Bryconamericus alpha

Branched analfin rays 17 or fewer; small red or yellowdot on upper caudal peduncle present Bryconamericus cismontanus


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Discussion

A comprehensive discussion of the relationships ofBryconamericus species from South America is notpossible at this time due to our poor knowledge of

the taxonomy and systematics of the genus. Nosynapomorphies presently define Bryconamericusas a monophyletic unit. However, in a recent studywith molecular characters RomnValencia &VanegasRos (in press) propose the first hypoth-esis for the monophyly of the Central Americanspecies of this genus. We include species assignedto the genus Knodusbecause we do not considera lack of scales on the caudal fin as sufficient towarrant generic recognition. Work in progresswill hopefully uncover osteological characters thatmay be useful for generic diagnosis, but at thistime the boundaries between AstyanaxBaird andGirard, Bryconamericus Eigenmann, Hemibrycon

Gnther, Hemigrammus Gill, HyphessobryconDurbin, and Moenkhausia Eigenmann, remaintenuous and arbitrary.

Bryconamericus singularis n. sp. is similar toBryconamericus orinocoense (RomnValencia,2003d, table 1), but can be distinguished by thelonger maxillary bone (18.332.9% in B. singularisvs. 12.831.4%), and the shape of the opercle,

which has a straight posterior edge vs. strong notchpresent in upper part of opercle. It also differs in thelower number of vertebrae (33 in B. singularis vs.3536), is longer and less deep bodied: maximumbody depth (19.525.9% en B. singularisvs. 26.831.5%), and has a longer distance separating thedorsal fin and the hypurals (46.956.2 % in B.singularis vs. 36.439.9%). The upper jaw is alsolonger: (19.931.1% in B. singularis vs. 11.717.8%); and the shape of the posterior edge of theopercle is concave and lacks a notch in B. singularis.

The presence of six to seven small unicuspidteeth in B. singularis coincides with the five toseven small teeth reported in B. turiubaLangeani et

al. (Upper Ro Paran system) by Langeani et al.(2005) but that species has tricuspid as well asunicuspid teeth in this series.

Key to the species of Bryconamericus from the Guyana Shield.

Clave para las especies de Bryconamericus del Macizo de Guyana.

1 Fewer than 34 lateral scales 2

More than 33 lateral scales 3

2 Unbranched dorsal fin rays usually ii; unbranchedanalfin rays usually iv; length of maxilla 3239%of head length, mean 35%; body short and stocky,its greatest body depth 2530% standard length,mean 27% Bryconamericus orinocoense

Unbranched dorsal fin rays usually iiiiv; unbranchedanalfin rays usually iii; length of maxilla in head length1832%, mean 27%; body elongate and slender,its greatest body depth 1926% standard length,mean 24% Bryconamericus singularis

3 Central caudalfin rays with a black stripe, or

with a dark spot on their bases 4Central caudalfin rays without black stripe,a vertically oriented, crescent-shaped blotch at base(sometimes diffuse) usually present Bryconamericus cinarucoense

4 More than 19 branched analfin rays Bryconamericus alpha

Fewer than 20 branched analfin rays 5

5 Lateral scales 4041; caudal fin overall dusky, withdarker spot on base of central caudal rays, and whitishspot at base of upper lobe, and sometimes aat base of lower lobe as well Bryconamericus macrophthalmus

Lateral scales 3239; caudal fin not pigmented as above 6

6 Caudal peduncle with small red or yellow dot;maxilla with 12 teeth with cusps of equal length;lateral scales 3238; branched analfin rays 1318 Bryconamericus cismontanus

Caudal peduncle lacks red or yellow dot; maxilla with46 teeth, with central cusp longer than others;lateral scales 3839; branched analfin rays 1718 Bryconamericus subtilisform


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While preparing the descriptions of B. singularis(in this work) and B. orinocoense (RomnValen-cia, 2003d) we noted that we could place them inthe genus Moenkhausiaaccording to some characterstates (Eigenmann, 1918). For example, both have

the teeth in the outer row of the premaxilla in astraight row and five teeth in the inner row. How-ever, both species lack scales on the base of thecaudal fin, as would be the case if they wereMoenkhausia(some Bryconamericus, however, alsohave a scaled caudal, and some authors placethem in the genus Knodus because of this). Thesecond infraorbital is not in contact with thepreopercle in Moenkhausiaspecies but is in contactin B. singularis and B. orinocoense. This situationsimply lends further credence to the paraphyly ofboth Bryconamericus and Moenkhausia as cur-rently conformed, and points out the need for abroad revision and redefinition of these, and most

other characid genera.Based on the results and analysis of this study

(for example, the reidentification of specimensfrom the localities where records of B. brevicepsyB. heterestheswere previously purported to occur,and the characters presented in the keys) we con-sider previous reports of B. breviceps and B.heteresthes from Venezuela (RomnValencia,2005) to be misidentifications, and they are hereconsidered as either B. cinarucoense, or another,as yet undescribed species.

On comparing data obtained during this studywith previous reports (RomnValencia, 2003d,2005) we can find no differences to substantiate therecognition of B. alpha and B. motatanensis asseparate species. In only one character: distancefrom dorsalfin origin to analfin origin, is there asmall difference: (35.836.1% in B. alphavs. 37.048.1% in B. motatanensis) which we considerinsufficient. Thus, based on the International Codeof Zoological Nomenclature (1999), the valid nameis B. alpha, and B. motatanensis (described as B.beta motatanensis) is considered a synonym.

Acknow ledgements

We thank Carmen Montaa, D. Aubrey Arrington, andCraig Layman who collected many specimens ofthese new species during their research on the ecol-ogy of the fishes of the Cinaruco River as part of theirthesis work under the supervision of Dr. KirkWinemiller. We also thank the Universidad NacionalExperimental de los Llanos Occidentales EzequielZamora'' for their support of our research, FONACITfor funds that support the Fish Collection of theMuseum of Zoology of UNELLEZGuanare, andINAPESCA for scientific collecting permits. We givespecial recognition to the personnel of the Fish Col-lection at UNELLEZGuanare: Keyla Marchetto,Luciano Martnez and Iraima Montaa for their help in

processing the specimens. This report was madepossible by the generous support of COLCIENCIAS,of the Universidad de Quindio, Facultad de Ciencias

Bsicas Programa de Biologa and Vicerrectora deinvestigaciones, for the trips made by C. RV. to themuseums in Guanare (MCNG), MaracayRanchoGrande (EBRG) and Caracas (MBUCV) in Venezuela.

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