Britski Birindelli 2016 Leporinus Altipinnis

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Ichthyological Exploration of FreshwatersAn international journal for field-orientated ichthyology

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Ichthyol. Explor. Freshwaters, Vol. 27, No. 1

25Copyright © Verlag Dr. Friedrich Pfeil

Ichthyol. Explor. Freshwaters, Vol. 27, No. 1, pp. 25-40, 7 figs., 1 tab., April 2016© 2016 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902

Redescription of Leporinus altipinnis ,a senior synonym of L. falcipinnis ,

and comments on L. holostictus (Characiformes: Anostomidae)

Heraldo A. Britski* and José L. O. Birindelli**

Leporinus altipinnis is redescribed based on type specimens and additional material from the Amazonas andOrinoco river basins in Brazil and Venezuela. It is distinguished from congeners by having a unique color patterncomposed by five dark transversal bars on the trunk in specimens up to 40 mm SL, which split into eight bars inspecimens ranging from 60 to 100 mm SL, then into ten bars in specimens of approximately 140 mm SL, and theninto 13 or 14 dark bars in specimens larger than 200 mm SL. Leporinus altipinnis is also the only species of banded Leporinus that shows anteriormost rays of dorsal and caudal fins elongate, giving them falcate aspect. Leporinusaltipinnis inhabits preferably black-water rivers, and is distributed in tributaries of the Amazonas river basin, suchas the Arapiuns and Negro rivers, and in the Orinoco river basin. Leporinus falcipinnis is recognized as a juniorsynonym of L. altipinnis. In addition, comments on L. holostictus are provided and the species is considered a

junior synonym of L. fasciatus.

Leporinus altipinnis é redescrita com base nos exemplares tipo e em exemplares adicionais das bacias dos riosAmazonas e Orinoco, no Brasil e Venezuela. A espécie se diferencia de suas congêneres pelo padrão de coloridoexclusivo, composto de cinco faixas transversais escuras no tronco nos exemplares até 40 mm SL, as quais sedividem em oito faixas nos exemplares de 60 a 100 mm SL, depois em dez faixas nos exemplares com aproxima-damente 140 mm SL, e em treze ou quatorze faixas nos exemplares com 200 mm SL ou mais. Leporinus altipinnis é também a única espécie do grupo com faixas transversais que apresenta prolongamento dos raios mais ante-riores das nadadeiras dorsal e caudal, dando-lhes aspecto falcado especialmente às nadadeiras dorsal e caudal.A espécie tem preferências por rios de águas pretas e ocorre em afluentes de rios amzônicos, especialmente nosrios Arapiuns e Negro e na bacia do rio Orenoco. Leporinus falcipinnis é aqui considerada um sinônimo júnior de

L. altipinnis. Adicionalmente, são fornecidos comentários sobre L. holostictus e a espécie é considerada um sinôni-mo júnior de L. fasciatus.

* Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42494, 04218-970, São Paulo, SP, Brazil.E-mail: [email protected]

** Departamento de Biologia Animal e Vegetal, Universidade Estadual de Londrina, Caixa Postal 10.011, 8657-090, Londrina, PR, Brazil. E-mail: [email protected] (corresponding author)


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Britski & Birindelli: Redescription of Leporinus altipinnis

Introduction

Leporinus is one of the largest genera in the or-der Characiformes, with approximately 80 validspecies (Garavello & Britski, 2003; Sidlauskas &

Vari, 2008). Recent collecting trips and increasinganalysis of specimens deposited in fish collectionshave revealed a still great number of undescribedspecies in the genus (e. g., Birindelli & Britski,2009, 2013), as well as many taxonomic problemsthat need to be investigated (e. g., Britski et al.,2011, 2012). Small individuals (15-30 mm SL) of all speciesof Leporinus present color patterns with transversedark brown bars on the body, usually in numberof eight. During development, this color pattern

changes in most species of the genus, with the dark bars fading and dark brown blotches graduallyappearing and even forming, in some cases, longi-tudinal dark brown stripes on the body (Birindelli& Britski, 2013). In some species of Leporinus, theoriginal dark bars may persist in adult individu-als, in which case, these bars are either presentas in juveniles or split partially acquiring Y-, V-,K- or X-shaped bars, or split completely into twoor more dark transversal bars increasing up to15 the total number of bars in fully-developedspecimens. All species in which dark transversal bars persist in fully developed specimens aretreated as belonging to the ‘ banded species group’of Leporinus, following Böhlke (1958), Garavello& Britski (1987), and Sidlauskas & Vari (2012).Sidlauskas & Vari (2008) recognized the bandedspecies of Leporinus as possibly a monophyleticunit supported by some osteological features,including the presence of nine branched pelvic-finrays (vs. eight in all other anostomids; Sidlauskas& Vari, 2008, character #111). The monophyly ofthe banded species of Leporinus was also sup-ported by molecular data (Ramirez et al., 2016).Böhlke (1958), who studied some species of thisgroup stated that “the various banded species ofLeporinus have never been adequately comparedwith a view toward determining how manyand which are the valid species”. Subsequently,few other authors studied the taxonomy of thisgroup (e. g., Géry, 1964, 1977, 1999). Sidlauskas &Vari (2012) studying the anostomid species fromGuyana highlighted the taxonomic confusion stillpersistent in many valid species of the banded

species of Leporinus.

The banded species of Leporinus is hereindefined as including all species having darktransversal bars in fully-developed specimensand nine branched pelvic-fin rays. According tothe most recent list of valid species (Garavello &

Britski, 2003), ten species are included into the banded species of Leporinus: L. affinis Günther,1864, L. bleheri Géry, 1999, L. desmotes Fowler,1914, L. falcipinnis Mahnert, Géry & Muller, 1997,L. fasciatus (Bloch, 1794), L. holostictus Cope, 1878, L. jatuncochi Ovchynnyk, 1971, L. pearsoni Fowler,1940, L. tigrinus Borodin, 1929, and L. yophorus Eigenmann, 1922. Note that L. octofasciatus Stein-dachner, 1915, previously considered as part ofthat group (e. g., Garavello, 1979), is excluded byhaving only eight branched pelvic-fin rays.

Through the study of specimens of distinctontogenetic stages of Leporinus deposited in differ-ent collections, one of the pieces of the taxonomicpuzzle involving the banded species of Leporinus was solved. The aim of the present contributionis to present the results obtained during thisinvestigation, which consist of the redescriptionof L. altipinnis based on type specimens and ad-ditional material, diagnosing it among congeners,and the recognition of L. falcipinnis as its juniorsynonym. In addition, details concerning the colorpattern of L. fasciatus, and color pattern shiftsduring ontogeny, are provided and the speciesL. holostictus is recognized as a junior synonymof L. fasciatus.

Materials and methods

This paper is part of a broader research projectthat aims to revise all banded species of Leporinus. For that, all nominal species of the banded spe-cies group of Leporinus were evaluated. Extensivematerial was examined from distinct fish collec-tions, including type specimens of all species,and partial or complete ontogenetic series ofmost species. Primary type specimens of L. affinis,L. desmotes, L. falcipinnis, L. holostictus, L. pearsoni,L. tigrinus, and L. yophorus were examined. Theholotypes of L. fasciatus (ZMB 3543, availablein Zarske, 2011) and L. jatuncochi (MUSM 5868)were examined through photographs. One of theparatypes of L. bleheri was examined (instead ofthe holotype). The limits between valid species

were determined through comparisons of exten-sive ontogenetic series with particular attention


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to color pattern shifts, and of meristic data withparticular attention to number of scales in lateralline and transversal series, and number of teethin the premaxillary and dentary bones. Counts and measurements were taken accord-

ing to Britski & Garavello (1978) and Winterbot-tom (1980). Standard length (SL) is expressed inmm and all other measurements are expressedas percentage of SL, except subunits of the head,which are expressed as percentages of headlength. The lateral-line scale count included thepored scales extending onto the base of the mediancaudal-fin rays; counts of the longitudinal scalerows above lateral line exclude the lateral-linescale and the middorsal scale row; counts of thelongitudinal scale rows below lateral line exclude

the lateral-line scale row and include half a scalerow when the pelvic-fin origin is immediately behind the middle of a scale. Weberian-apparatusvertebrae were counted as four elements andincluded in the vertebral counts; the compoundcaudal centrum (i. e., preural 1 fused to ural 1)of the caudal region was counted as a single ele-ment. The pattern of radii was defined on scalesimmediately dorsal to the lateral line row at thevertical through the dorsal-fin origin. All examined specimens are alcohol pre-served, except when indicated by CS, clear andstained according to Taylor & Van Dyke (1985),or SK, dry skeleton prepared following gener-ally the methods of Bemis et al. (2004). Institu-tional abbreviations are: ANSP, Academy ofNatural Sciences of Philadelphia, Philadelphia;BMNH, Natural History Museum, London; CAS,California Academy of Sciences, San Francisco;FMNH, Field Museum of Natural History, Chi-cago; INPA, Instituto Nacional de Pesquisas daAmazônia, Manaus; MBUCV, Museo de Biologíade la Universidad Central de Venezuela, Caracas;MCNG, Museo de Ciencias Naturales de Guanare,Guanare; MCP, Museu de Ciência e Tecnologiada PUCRS, Porto Alegre; MCZ, Museum ofComparative Zoology, Boston; MNRJ, MuseuNacional do Rio de Janeiro, UFRJ, Rio de Janeiro;MPEG, Museu Paraense ‘Emílio Goeldi’, Belém;MZUEL, Museu de Zoologia da UniversidadeEstadual de Londrina, Londrina; MZUSP, Museude Zoologia da Universidade de São Paulo, SãoPaulo; and UF, University of Florida, FloridaMuseum of Natural History, Gainesville.

Leporinus altipinnis Borodin, 1929(Figs. 1-2)

Leporinus fasciatus altipinnis Borodin, 1929: 280,pl. 8 (type locality: Jaturana; 2 syntypes MCZ

20487). – Fowler, 1950: 232, fig. 269 (catalogue,figure copied). – Mahnert, Géry & Muller,1997: 843 (reference).

Leporinus falcipinnis Mahnert, Géry & Muller, 1997:838 (type locality: rio Arapiuns, 1 hour aboutdownstream of mouth of rio Aruá, small farmat right shore, lower Tapajós basin, Pará,Brazil, 2°04' S 55°38' W; holotype MZUSP51827). – Garavello & Britski, 2003: 75 (cata-logue). – Britski & Garavello, 2007: 25 (cata-logue). – Sidlauskas & Vari, 2008: 73, 150, 152,

179 (Venezuela, Amazon, backwater of islandin Rio Atabapo, about 1 hour above SanFernando de Atabapo; some anatomical andmorphological data). – Santos et al., 2006: 36,fig. (illustration, diagnosis, natural history,fishery).

Leporinus sp. – Toledo-Piza, 2002: 103, drawingnumber 118 (Venezuela, San Carlos, rio Negro,illustration).

Diagnosis. Leporinus altipinnis is distinguishedfrom all congeners by having a unique colorationcomposed by five dark brown transversal bars onthe trunk of specimens up to 40 mm SL, whichsplit into eight bars in specimens 60-100 mmSL, then into ten bars in specimens of approxi-mately 140 mm SL, and at last into 13 or 14 barsin specimens larger than 200 mm SL (Figs. 1-2).Leporinus altipinnis is further distinguished fromall congeners, except L. affinis, L. bleheri, L. des-motes, L. fasciatus, L. jatuncochi, L. octofasciatus,L. pearsoni, L. tigrinus, and L. yophorus, by hav-ing dark transversal bars on body in adults (vs. body with one to several dark round blotches, orwith one to six dark stripes) and nine branchedpelvic-fin rays (vs. eight). Leporinus altipinnis is distinguished from L. affinis, L. bleheri, L. des-motes, L. jatuncochi, L. octofasciatus, L. pearsoni, L. tigrinus, and L. yophorus by having 10-14 darktransversal bars in specimens larger than 140 mmSL (vs. 6-8); and anteriormost rays of the dorsaland caudal fins elongated, distinctly longer thanmedian rays in specimens larger than 100 mm SL(vs. anterior rays of dorsal and caudal fins not

distinctly longer than adjacent rays in specimenslarger than 100 mm SL). Leporinus altipinnis isdistinguished from L. fasciatus by having 6 or 7


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scale rows between lateral line and dorsal-finorigin (vs. 7 or 8), and 13 to 14 bars on body inspecimens larger than 140 mm SL (vs. 9 or 10).

Description. Morphometric data in Table 1.

Medium size, relative to congeners. Largest ex-amined specimen 315.7 mm SL. Head and bodyelongate and moderately compressed. Dorsalprofile straight from snout tip to supraoccipitalspine, sometimes slightly concave between an-terior and posterior nares, gently convex fromsupraoccipital to dorsal-fin origin; straight alongdorsal-fin base, straight or gently convex fromterminus of dorsal-fin base to adipose-fin origin,distinctly convex from adipose-fin origin to baseof anteriormost procurrent rays of caudal fin. Ven-

tral profile convex from lower jaw tip to verticalthrough posterior naris, straight from that pointto vertical through pectoral-fin base, straight toslightly convex from latter point to anal-fin origin,straight along anal-fin base, and concave fromterminus of anal-fin base to base of anteriormostprocurrent rays of caudal fin. Greatest body depthat dorsal-fin origin. Mouth terminal, mouth opening longitudinal-ly aligned with ventral margin of pupil in speci-mens of 200 mm SL or larger, and longitudinallyaligned with dorsal margin of pupil in specimens50-100 mm SL (Figs. 1-2). Snout rounded, anteri-orly blunt. Premaxilla with four incisiform teethgradually decreasing in size from symphysealtooth. Dentary with four incisiform teeth gradu-

ally decreasing in size posterolaterally (Fig. 3). Scales cycloid, with 5 (3) or 6 (4) radii. Lateralline complete, extending from supracleithrum to base of median caudal-fin rays, with 41 (1), 42 (16)or 43 (19) pored scales (42 on syntypes of L. altipin-

nis, 43 on holotype of L. falcipinnis). Longitudinalscale rows between dorsal-fin origin and lateralline 6 (32) or 7 (3) (6 on syntypes of L. altipinnis,7 on holotype of L. falcipinnis). Longitudinal scalerows between lateral line and pelvic-fin origin5 (15) or 6 (21) (6 on syntypes of L. altipinnis, andholotype of L. falcipinnis). Longitudinal scale rowsaround caudal peduncle 16 (36). Predorsal scalesfrom tip of supraoccipital spine to dorsal-fin origin12 (7), 13 (11), 14 (10), 15 (2) or 16 (1). Dorsal-fin rays ii,10 (31). Dorsal-fin origin

slightly anterior to midpoint of standard lengthand to vertical through pelvic-fin origin; distalmargin of dorsal fin strongly concave in speci-mens of 150 mm SL or larger, and slightly con-vex in specimens of 100 mm SL or smaller, lastunbranched and first two or three branched rayselongate in most specimens larger than 100 mmSL. Adipose fin small, teardrop-shaped withorigin approximately at vertical through base oflast anal-fin ray. Pectoral-fin rays i,13 (1), i,14 (14),i,15 (15), or i,16 (1) (13 on syntypes of L. altipinnis,16 on holotype of L. falcipinnis); tip of adpressedpectoral fin extending to midpoint betweenorigins of pectoral and pelvic fins or up to fifth,fourth or third scale anterior to pelvic-fin base;distal margin of pectoral fin concave posteriorly

Table 1. Morphometric data for Leporinus altipinnis. N, number of specimens; SD, standard deviation; ALT, syn-types of L. fasciatus altipinnis, MCZ 20487; FAL, holotype of L. falcipinnis, MZUSP 51827.

n mean range SD ALT ALT FAL

Standard length (SL) 35 53.0-289.0 92.0 80.0 223.0

Percents of standard lengthPredorsal distance 31 47.1 45.1-51.5 1.3 48 48 46Dorsal-fin to adipose-fin 31 38.9 35.7-40.8 1.2 40 39 40Prepelvic distance 31 49.0 45.9-52.2 1.4 50 49 46Body depth 31 24.7 23.0-26.8 0.9 25 26 25Caudal peduncle depth 31 9.2 8.2-9.9 0.4 10 10 10Caudal peduncle length 31 12.7 11.4-13.7 0.5 12 13 13Anal-fin lobe length 31 14.7 13.8-15.7 0.5 14 14 15Head length 31 24.6 22.0-26.4 1.1 25 26 22

Percents of head lengthPreopercle length 31 78.4 74-82 1.9 79 77 74Snout length 31 37.4 31-42 2.4 36 39 41

Head depth 31 75.0 69-

84 3.3 73 75 82Eye diameter 31 28.3 19-35 3.9 29 27 20Bony interorbital 31 37.3 34-43 2.3 38 35 43



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Fig. 1. Leporinus altipinnis: a-b, syntypes of L. fasciatus altipinnis, MCZ 20487, 92.0 and 80.0 mm SL; Brazil: Jatu-arana, Amazonas basin; c, holotype of L. falcipinnis, MZUSP 51827, 223.0 mm SL; Brazil: rio Arapiuns, Tapajós

basin.

a

b

c

and gently convex ventrally, unbranched andfirst two or three branched rays slightly elon-gate in most specimens larger than 100 mm SL.Pelvic-fin rays i,9 (31); distal margin of pelvic finslightly concave posteriorly and slightly convexmedially, unbranched and first two or three branched rays elongate in most specimens largerthan 100 mm SL. Anal-fin rays ii,7 (1), ii,8 (29), orii,9 (1); anal-fin origin approximately at verticalthrough posterior margin of fourth or fifth scaleanterior to adipose-fin origin; distal margin ofanal fin concave in most examined specimenslarger than 100 mm SL, last unbranched and firsttwo or three branched rays elongate in most of

those specimens; adpressed anal fin not reaching base of caudal-fin rays. Principal caudal-fin raysi,8,9,i (31). Caudal fin forked, unbranched and first

two or three branched rays of each lobe elongatein most specimens, making fin lobes distinctlyfalcate. Supraneurals 5 (2). Vertebrae 40 (2), withribs on vertebrae five through 20 (2).

Color in alcohol. Ground color brown or tan,weakly countershaded. Body with conspicuoustransverse dark brown bars generally wide andencircling head and trunk, including ventralregion. Head with broad dark brown bar overdorsal portion of snout tip, skin covering upper jaw, and lower jaw ventrally; broad dark brown bar on dorsal area between eyes continued ven-trally below eye. Trunk with five brown transverse

bars in specimens up to 40 mm SL, which splitinto eight bars in specimens ranging from 60 to100 mm SL, then into ten bars in specimens of


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Fig. 2. Leporinus altipinnis: a, MZUSP 51011, 36.0 mm SL; b, MZUSP 29139, 95.7 mm SL; c, MZUSP 91583, 229.8 mmSL; d, MZUEL 7052, 131.1 mm SL; e, uncatalogued specimen, approximately 200 mm SL. All from Brazil: rioNegro, Amazonas basin (except d, aquarium trade specimen); c-d photographed alive.

a

b

c

d

e



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approximately 140 mm SL, and at last into 13 or14 dark bars in specimens larger than 200 mm SL(Fig. 2). Membranes of fins generally hyaline, rayssomewhat darkened in specimens of 150 mm SLor larger, brown blotches at base of dorsal and

anal fins and at middle of pelvic fin in specimensof approximately 50 mm SL, blotches continuouswith dark bars of body. Adipose fin with base anddistal margin dark brown.

Color in life. Life coloration similar to preservedspecimens, but ground coloration more yellowand fins more hyaline (Fig. 2d-e).

Distribution. Leporinus altipinnis inhabits prefer-ably black-water rivers, and is known from rio

Arapiuns (a tributary of the lower rio Tapajós),a small creek near Parintins (type locality), rioParaconi at FLONA do Pau Rosa, and from sev-eral localities in the rio Negro and río Orinoco,in Brazil and Venezuela (Fig. 4). There is no sig-nificant geographical variation in meristic dataor coloration of the specimens from the Amazonand Orinoco basins, what may be due to fact thatall examined samples came from similar habitats(i. e., blackwater rivers of lowlands of the rain for-est) or because the Cassiquiari canal is connectingpopulations from those two basins.

Remarks. Borodin (1929: 280) described Leporinusaltipinnis on the basis of two specimens collected by the Thayer Expedition and referred to the typelocality as ‘ Jaturana, Brazil’. This name, however,is absent from the list of localities of the ThayerExpedition (Dick, 1977; Higuchi, 1996). There is,on the other hand, one locality recorded in thatlist similar to the one referred by Borodin (1929),‘ Jatuarana’ (Higuchi, 1996). Based on that, it islikely that the type locality was misspelled byBorodin. Higuchi (1996) indicate that Jatuaranarefers to a creek or small lake SW of Parintins, nearBarreirinha (2°47' S 57°04' W), Amazonas state,with specimens being collected by Mr. Naves.The latter was not a participant of the ThayerExpedition and was also credited as collectorof specimens labeled as collected in the ‘LagoHyanuary’, referred as Paraná do Januari, at ap-proximately 3°12' S 60°05' W, near Manaus, by Hi-guchi (1996). The collections credited to Mr. Naveswere obtained on January 10 and 15, 1866, when

the Thayer Expedition was in Manaus (Agassiz& Agassiz, 1868). Since specimens of L. altipinnis are known to be sold in the fish market at Manaus

(Santos et al., 2006), it is also possible that the typespecimens were purchased in Manaus from Mr.Naves, who might have collected the specimensaround Manaus. A third possibility includes anarea in Brazil called ‘Floresta Nacional de Jatu-arana’ in Borba, Amazonas state, which has as boundary the rio Sucunduri, a tributary of the rioMadeira, approximately at 7°35' S 59°31' W. Eventhough the type locality is uncertain at this point,we follow Higuchi (1996) in considering Parintinsas the type locality of L. altipinnis. The naturalist Alfred R. Wallace draw a speci-men about 300 mm SL collected by him in the rioNegro in 1850-1852, and identified 150 years lateras Leporinus sp. (Toledo-Piza, 2002: 103, drawingnumber 118). The specimen displays 13 dark barson the body, a hallmark of L. altipinnis. Wallacealso recorded a few meristic data of the specimen,including 10 total rays in the pelvic fin (probablynine branched), a feature exclusive to the bandedspecies Leporinus, as mentioned above. It is in-teresting to note that the species was describedapproximately 80 years later, and it was until nowpoorly understood. Leporinus altipinnis has been labeled in muse-um collections from Venezuela (MCNG, MBUCV),in some publications (e. g., Montaña et al., 2010),and on websites as ‘Leporinus cf. fasciatus var.doble raya’, ‘mije doble ray’, etc., in reference tothe split of the dark bars during development.

Santos et al. (2006: 36) presented a drawing ofa large specimen of L. altipinnis bearing 15 dark bars on body. Even though, the largest specimens

Fig. 3. Leporinus altipinnis, MZUSP 110814, 228.0 mm

SL, head, showing jaws and teeth morphology. Brazil:rio Negro, Amazonas basin. Scale bar 10 mm.

angulo-articular

maxilla

premaxilla

retroarticular

dentary

angulo-articular

maxilla

premaxilla

retroarticular

dentary


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examined by us had only 13 or 14 dark bars,the second bar is dorsally split indicating thatspecimens larger than the ones herein examinedmight have 15 dark bars on body. In fact, Santoset al. (2006) mentioned that the species reachesup to 350 mm SL, whereas the largest specimenexamined herein is 315.7 mm SL. The only records of live specimens of L. altipin-nis includes Roggo (1996: 55, fig. 2), mentioned

by Mahnert et al. (1997) as a photograph of a livespecimen taken underwater in the rio Tapajós;Roggo (2015: photo F62-04, in ‘The freshwaterproject in the Rio Negro and Tributaries’), whichshows a group of five specimens of approximately100 mm SL with eight dark bars on the body;and Willis (2005), which includes a picture of aspecimen collected in the río Pasiba, a tributary

of the río Cassiquiare and identified as Leporinus cf. fasciatus var. ‘doble raya’.

Discussion

Borodin (1929: 280) described Leporinus altipinnis based on two specimens (“95 and 80 mm long”)with ten dark bars on the body. Borodin’s descrip-tion is short, possibly because he allocated thenew taxon as a subspecies of L. fasciatus, but the

accompanying drawings reveal precise featuresof coloration, position of the mouth, and shapeand arrangement of the teeth. On the other hand,

Mahnert et al. (1997) described L. falcipinnis onthe basis of a single large individual (220 mmSL) that possessed the maximum number of dark bars known for the species (i. e., 14). Interestingly, both aforementioned taxa were named in allusionto the elongated fin lobes, an exclusive feature ofthis species. The lack of information regarding theontogeny of the coloration of the banded speciesof Leporinus led to the description of the samespecies twice. In addition, both descriptions were based on few specimens. Mahnert et al. (1997: 837)noted the limitation of describing a species basedon limited material: “The new species describedhere is therefore based on the unique preservedspecimen. We are conscious that such a descrip-tion is fragmentary, as no osteological detailscould be studied, the juvenile color pattern and,

mainly, the sexual dimorphism in this speciesremain unknown”. This taxonomic history, aswell as the comments cited above, emphasizesthe need for more comprehensive taxonomic revi-sions based on extensive material and consideringontogenetic changes, especially for large speciesof Anostomidae that undergo impressive changesduring development.

Eigenmann (1912: 308) was the first author todescribe the ontogenetic change in the colorationof the banded species of Leporinus, through the

examination of 63 specimens of L. fasciatus col-lected in the Essequibo River, and ranging from52 to 335 mm SL. He described these changes as:

Fig. 4. Northern portion of South America, showing the distribution of Leporinus altipinnis. , type locality ofL. fasciatus altipinnis; , type locality of L. falcipinnis.

-75°-80°

10°

-5°

-10°

5°

0°

-70° -65° -60° -55° -50°



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“Young with the snout, a bar through and overthe eye, and another across the opercle and nape, black; five bars encircling the body, one in frontof the dorsal, one below and behind the dorsal,one in front and one behind the anal, and one at

the end of the caudal peduncle. The first two splitinto two bars when a length of 75 mm is reached,the next two begin to split at length of 200 mm, thecomponents moving apart; in the adult there areten obscure bars about equally spaced, beginningwith the one across the opercle”. Some details inEigenmann’s description were not corroborated by our investigation, especially regarding the tim-ing of the splits (see observations below). Böhlke(1958) presented the changes in the colorationof L. pearsoni in specimens 90-163 mm SL. Géry

(1964), through the examination of specimensof L. fasciatus collected in the upper Amazonin Loreto, Peru, reported that specimens of ap-proximately 35 mm SL exhibits eight dark bars onthe trunk and specimens of about 90 mm SL tendark bars, which occurred due to splitting of thesixth and seventh dark bars during development,observations corroborated herein. Specimens of L. fasciatus from 13.8 to around20 mm SL have eight dark bars on body (Fig. 5a),not considering those on head: the first one is thenape bar, located at the region of body immedi-ately posterior to occiput dorsally and laterallycovering opercle; the second and third bars are between the nape bar and origin of dorsal fin;the fourth bar is located ventral to middle ofdorsal-fin base; the fifth bar is located betweenthe end of dorsal-fin base and vertical throughanus; the sixth bar is at anal-fin origin, anteriorto adipose-fin origin; the seventh bar is located below rear of adipose fin; and the eighth bar atend of caudal peduncle partially covering baseof caudal-fin rays. In some individuals up to18 mm SL (MZUSP 29148, 29150, 53639, 112210),the second and third bars and/or the fourth andfifth are so close that they are almost completelycontinuous (Fig. 5a). These bars might be closetogether and partially united forming X, H, Kor V in specimens 18.5-35.6 mm SL (Fig. 5a- b)(MZUSP 29150, 112210, 112213). Eight distinctlyseparated (and undivided) bars are present inspecimens 23.8-35.0 mm SL (Fig. 5b-c) (MZUSP29150, 48379). The sixth bar might be only partiallydivided and shaped as X, H, K or V in specimens

42.5-

300 mm SL (Figs. 5d, 6a-

c) (MCP 46072;MZUEL 10196; MZUSP 29125, 29151, 43444),or completely divided into two separate bars in

specimens 60.1-300 mm SL (Figs. 5e, 6d) (e. g.,MZUEL 10187, MZUSP 29125). The seventh baris partially divided in specimens 42.5-81.9 mmSL (MCP 21033; MZUSP 29124, 29125, 48384),and completely divided in specimens larger

than those. The division of the sixth bar usuallystarts from dorsal portion of body, whereas theseventh bar usually divides along its entire depth(Figs. 5d, 6a). In most specimens there is a totalof ten dark bars on the body: three from head todorsal-fin origin, one below dorsal-fin base, threefrom dorsal-fin terminus to anal-fin origin, one below adipose fin and two on caudal peduncle(Figs. 5e, 6d). Nevertheless, in some specimensthe sixth bar is never completely divided and isshaped as X, V, K or H (Fig. 6b-c) (MCP 46072,

MZUEL 10196, CAS 20168, FMNH 93087). In thelatter specimens, there are a maximum of ninedark brown bars on body (Fig. 6b-c). In addi-tion, in some large specimens with ten bars, thesecond, third, fourth and/or fifth bars might beX or Y shaped (Bloch, 1794; Zarske, 2011) (ANSP176829, MPEG 17503).

Leporinus holostictus , a junior synonym ofL. fasciatus. Leporinus holostictus was described by Cope (1878: 690) on the basis of a singlespecimen of 104 mm SL collected in the PeruvianAmazon, precise locality unknown (Fig. 7a). Thespecies was later redescribed by Fowler (1906:330) on the basis of the same specimen, whichwas illustrated, conspicuously showing eightdark bars on the trunk, and an additional speci-men briefly mentioned at the end of the speciesdescription “The other example shows the depthas 3 1/2” (Fig. 7b). Géry (1964, 1977) consideredL. holostictus as a subspecies of L. fasciatus, albeitstating that it was probably invalid. Garavello(1978) came to similar conclusion and includedL. holostictus as a junior synonym of L. fasciatus.Nevertheless, L. holostictus was later consideredvalid in Garavello & Britski (2003). The holotypeof L. holostictus has the same color pattern as speci-mens of L. fasciatus of approximately 50-150 mmSL, four teeth on premaxilla, four teeth on dentary,lateral line with 42 perforated scales, seven rowsof scales from the dorsal-fin to the lateral line, fiverows of scales from the lateral line to the pelvic-fin base, and 16 circumpeduncular scales. The otherspecimen (ANSP 21468) later described by Fowler

(1906) and possibly collected with the holotype ofL. holostictus, although badly preserved and dif-ficult to determine, has four teeth on premaxilla,


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Fig. 5. Leporinus fasciatus: a-

c, MZUSP 29150, Brazil: rio Tefé, Amazonas basin (a, 18.5 mm SL; b, 23.8 mm SL;c, 35.0 mm SL); d, MZUSP 48384, 73.1 mm SL; Brazil: Lago Janauacá, Amazonas basin; e, MZUSP 7641, 114.4 mmSL; Brazil: Lago José-Açu, Amazonas basin.

a

b

c

d

e



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Fig. 6. Leporinus fasciatus: a, uncatalogued specimen, approximately 80 mm SL; Venezuela: rio Ventuari, Orinoco basin; b, MCP 46072, 119.7 mm SL; Brazil: rio Jauaperi, Negro basin; c, MZUEL 10196, 174.1 mm SL; Brazil: rio Pit-inga, Uatumã basin; d, MZUEL 10187, 205.8 mm SL; Brazil: Igarapé do Boto, Uatumã basin. All photographed alive.

a

b

c

d


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four teeth on dentary, 43 perforated scales, sevenrows of scales from the dorsal-fin to the lateral

line, five rows of scales from the lateral line to thepelvic-fin base, and 16 circumpeduncular scales.Those data place the holotype of L. holostictus andthe aforementioned additional specimen entirelywithin the range presented by L. fasciatus. In addi-tion, the examination of extensive material fromPeru, and the study of the changes in coloration ofL. fasciatus during ontogeny (herein described andillustrated), leaves no doubts that the L. holostictus is a junior synonym of L. fasciatus.

Material examined. All specimens alcohol preservedunless noted. Leporinus affinis: BRAZIL: BMNH1849.11.8.52-53, 2 syntypes, 148.8-175.4 mm SL; Pará:rio Capim. – MCP 18214, 3, 100.0-109.0 mm SL; Goiás:Luís Alves, lakes of rio Araguaia. – MNRJ 25724, 1,150.6 mm SL; Mato Grosso: rio das Mortes, tributary ofrio Araguaia. – MZUSP 47828, 3, 185.0-262.0 mm SL;Goiás, Cocalinho: Lago Rico, rio Araguaia. – MZUSP48038, 1 CS, 85.0 mm SL; Pará: Jatobal: rio Tocantins,4°34' S 9°39' W. L. altipinnis: BRAZIL: MCZ 20487, 2 syntypes, 80.0-92.0 mm SL; Amazonas: Jatuarana [probably a creek orsmall lake near Barreirinha, Parintins, Amazonas, ap-proximately 2°38' S 56°43' W]. – MZUSP 51827, holotypeof L. falcipinnis, 223.0 mm SL; Pará: rio Arapiuns, tribu-tary of the lower rio Tapajós. – CAS 20169, 1, 185.9 mm

SL; Amazonas: rio Negro, at Castanheiro. – CAS 70567,1, 180.4 mm SL; Amazonas: Boa Futura: rio Xeruiuni.– FMNH 78805, 1, 168.4 mm SL; Amazonas: Manaus.

– MPEG 305, 2, 58.7-

62.1 mm SL; Amazonas: rio Negroat Ilha de Tamaquaré. – MPEG 312, 2, 35.9-141.5 mmSL; Amazonas: Barcelos: rio Negro at Lago da Ilha. –MPEG 691, 2, 138.5-302.0 mm SL; Amazonas: rio Negroat Anavilhanas. – MPEG 16120, 1, 153.8 mm SL; Ama-zonas: Maués: rio Paraconi, at FLONA do Pau Rosa,3°48'58" S 8°17'09.4" W. – MPEG 16147, 1, 134.5 mm SL;Amazonas: Maués: rio Paraconi, at FLONA do PauRosa, 3°49'49.7" S 58°17'26.5" W. – MZUEL 7052, 1,131.1 mm SL; purchased in aquarium store in SãoPaulo city. – MZUSP 6821, 3, 53.6-65.1 mm SL; Ama-zonas: Manaus: creek tributary of rio Tarumãzinho,

3°10' S 60°00' W. – MZUSP 29134, 11, 65.9-

83.0 mm SL;Amazonas: rio Arirará, rio Negro basin, 0°31' S 63°33' W.– MZUSP 29135, 4, 69.5-11.4 mm SL; Amazonas: Bar-celos: rio Negro, 0°58' S 62°57' W. – MZUSP 29136, 15,54.6-84.5 mm SL; Amazonas: rio Negro at Ilha de Cu-muru. – MZUSP 29138, 46, 53.0-87.5 mm SL; Amazonas:rio Negro at Ilha de Tamaquaré, 0°30' S 64°55' W. –MZUSP 29139, 41, 64.0-95.7 mm SL, 1 CS, 81.3 mm SL;Amazonas: rio Negro at Ilha de Buiu-Açu. – MZUSP29140, 1, 92.3 mm SL; Amazonas: rio Negro, below rioDaraá, 0°28' S 64°46' W. – MZUSP 29146, 3, 51.5-74.5 mmSL; Amazonas: Santa Isabel do Rio Negro: rio Negro atmouth of rio Urubaxi, 0°31' S 64°50' W. – MZUSP 31067,

1, 61.6 mm SL; Amazonas: rio Negro at Ilha de Tama-quaré, 0°30' S 64°55' W. – MZUSP 51011, 9, 36.0-51.0 mmSL; Amazonas: Manaus: rio Negro, 3°05' S 60°13' W. –

Fig. 7. Leporinus fasciatus: a, holotype of L. holostictus, ANSP 21467, 101.1 mm SL; unknown locality in the Peru-vian Amazon; b, ANSP 21468, 72.5 mm SL, possibly collected with holotype of L. holostictus.


a

b


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MZUSP 63461, 6, 48.1-65.4 mm SL; Amazonas: Parica-tuba: rio Negro marginal lagoon. – MZUSP 91437, 1,315.7 mm SL; Amazonas: Lago do rio Aiuanã, rio Negro

basin, 0°38' S 64°56' W. – MZUSP 91583, 2, 224.4-229.8 mm SL; Amazonas: rio Uaupés, tributary of rioNegro, 2°55' S 69°38' W. – MZUSP 110814, 2 SK, 220.0-

228.0 mm SL; Roraima: Caracaraí: rio Jufari, 1°01'46" S62°06'06" W. – MZUSP 112215, 1, 248.2 mm SL; Amazo-nas: Manaus: rio Negro, 3°10' S 60°00' W. – MZUSP112216, 1, 240.6 mm SL; Amazonas: Manaus: tributaryof igarapé Tarumãzinho, rio Negro basin. VENEZUELA:FMNH 104000, 1, 182.5 mm SL; Amazonas: San Fer-nando de Atabapo: río Atabapo. – FMNH 104017, 1,289.0 mm SL; Amazonas: río Autana, 4°43'42" N 67°38'07" W. – MCNG 21378, 2, 160.9-161.2 mm SL; AP:Laguna Larga, río Cinaruco. – MCNG 3297, 8, 197.9-199 mm SL; Amazonas: río Marapiare, in front ofYutaje. – MCNG 41968, 3, 50.4-67.4 mm SL; Dpto. Río

Negro: Laguna Anapacoa at Piedra Anapacoa, ríoPasimoni, 1°50'81" N 66°35'11" W. Leporinus bleheri: BRAZIL: MZUSP 58375, 1 paratype,114.5 mm SL; Bolivia: Itenéz: río Verde, tributary of ríoGuapore, 14°08' S 60°30' W. – MPEG 10791, 4, 128.1-154.3 mm SL; Amazonas: Coari. – MPEG 16764, 2,169.0-180.6 mm SL; Amazonas: Coari. – MZUSP 66676,1, 130.0 mm SL; Amazonas: Santa Isabel do Rio Negro,rio Tiquié, 0°16'27" N 69°54'56" W. – MZUSP 85374, 1,168.3 mm SL; Amazonas: Santa Isabel do Rio Negro:rio Tiquié, 0°15'22" S 69°50'23" W. – MZUSP 93445, 1,168.0 mm SL; Amazonas: Santa Isabel do Rio Negro:rio Tiquié, 0°10'N 69°07' W.

L. desmotes: GUYANA: ANSP 39324, holotype,146.4 mm SL; ANSP 39325, 3 paratypes, 84.4-121.3 mmSL; Rupununi river, paratypes. – BMNH 1972.7.27.124,1, 69.5 mm SL; Guyana: Arakwai creek, Rupununi basin.BRAZIL: INPA 2666, 1, 98.6 mm SL; Roraima: rio Tacutu.– INPA 4611, 5, 117.7-171.4 mm SL; INPA 15431, 4,82.4-140.3 mm SL; Amazonas: Uraricoera: rio Branco.– MZUSP 40734, 2 alc, 119.0-127.6 mm SL; Goiás:Monte Alegre de Goiás: rio Paranã, tributary of rioTocantins. – MZUSP 47906, 1, 125.3 mm SL; Pará: rioCupijó, tributary of rio Tocantins. – MZUSP 105499, 1,126.8 mm SL; Pará: Marabá: rio Tapirapé, 5°36'47" S50°26'42" W. – MZUSP 107802, 1, 106.2 mm SL; Pará:Canaã dos Carajás: rio Paranapanema, at rapids belowPoço do Jaú, 6°23'15" S 50°02'03" W. L. fasciatus: BRAZIL: CAS 20167, 3, 104.3-200.0 mmSL; Amazonas: rio Negro at Castanheiro. – CAS 139291,1, 243.0 mm SL; Amazonas: rio Amazonas at VillaBella. – MCP 21033, 1, 81.9 mm SL; Pará: Santarém: rioTapajós at Lago Verde, 2°31' S 54°57' W. – MCP 30008,2, 90.9-96.3 mm SL; Amazonas: Alvarães: Lago Amanã,at mouth of rio Baré, 2°27' S 64°43' W. – MCP 46072, 1,119.7 mm SL; Roraima: Rorainópolis: rio Jauaperi,0°13'55" S 61°03'52" W. – MZUEL 10187, 1, 205.8 mm SL;Amazonas: Presidente Figueiredo: Igarapé do Boto,

tributary of rio Uatumã, 2°07'00" S 59°18'24" W. – MZUEL10194, 2, 228.0-242.2 mm SL; Amazonas: PresidenteFigueiredo, rio Uatumã at Cachoeira Morena, 2°07'21" S

59°34'43" W. – MZUEL 10196, 13, 150.4-204.5 mm SL;Amazonas: Presidente Figueiredo: rio Pitinga, tributaryof rio Uatumã, at Corredeira Quarenta Ilhas, 0°53'16" S59°34'28" W. – MZUEL 11547, 2, 114.5-117.6 mm SL;Amazonas: Presidente Prudente: rio Pitinga, tributaryof rio Uatumã, 1°06'46" S 59°36'26" W. – MZUSP 3581,

3, 132.5-

173.4 mm SL; Pará: Santarém: rio Tapajós. –MZUSP 3816, 2, 217.2-221.5 mm SL; “Pará and Ama-zonas”. – MZUSP 5428, 10, 178.0-225.0 mm SL; Pará:Oriximiná: rio Trombetas. – MZUSP 5808, 1, 288.0 mmSL; Amazonas: Silves: Lago Saracá. – MZUSP 5883, 1,228.7 mm SL; Amazonas: Manacapuru: Lago Manaca-puru. – MZUSP 6071, 2, 178.1-198.5 mm SL; Amazonas:Manaus: rio Preto da Eva. – MZUSP 6095, 1, 154.5 mmSL; Amazonas: rio Puraquequara, 2°56' S 59°49' W. –MZUSP 6146, 1, 266.5 mm SL; Amazonas: Manaus: rioNegro, 3°10' S 60°00' W. – MZUSP 6186, 1, 250.0 mm SL;Amazonas: Manaus: Igarapé Jaraqui, tributary of rio

Negro. – MZUSP 6770, 3, 93.9-

253.0 mm SL; Amazonas:Manaus: tributary of Igarapé Tarumãzinho, rio Negro basin. – MZUSP 7055, 1, 160.7 mm SL; Amazonas: rioCanumã, rio Madeira basin, 4°58' S 58°56' W. – MZUSP7641, 1, 114.4 mm SL; Amazonas: Parintins: Lago José-Açu. – MZUSP 13209, 1, 104.9 mm SL; Amazonas: RioPreto da Eva: 2°42' S 59°42' W. – MZUSP 13211, 11,72.5-293.0 mm SL; Pará: rio Trombetas at Lago Jacaré.– MZUSP 13456, 1, 202.8 mm SL; Amazonas: Itacoatiara:rio Amazonas. – MZUSP 13902-4, 3, 247.0-267.5 mmSL; Rondônia: rio Machado at Lago do Paraíso. – MZUSP15680, 2, 222.7-244.3 mm SL; Pará: Oriximiná: rio Trom-

betas, 1°25' S 56°37' W. – MZUSP 15725, 1, 290.5 mm SL;

Pará: rio Trombetas. – MZUSP 15819, 3, 172.2-

238.0 mmSL; Pará: rio Trombetas at Reserva Biológica do RioTrombetas. – MZUSP 27387, 1, 214.0 mm SL; Amazonas:rio Negro at Anavilhanas. – MZUSP 27901, 2, 60.4-88.7 mm SL; Amazonas: Iranduba: Lago Janauacá, 3°13' S60°04' W. – MZUSP 29125, 30, 30.8-111.5 mm SL, 2 CS,97.1-105.5 mm SL; Amazonas: rio Negro, below rioDaraá, 0°28' S 64°46' W. – MZUSP 29126, 1, 123.8 mmSL; MZUSP 29127, 3, 52.8-112.0 mm SL; MZUSP 29128,1, 103.2 mm SL; MZUSP 29129, 1, 112.4 mm SL; MZUSP29132, 1, 54.2 mm SL; Amazonas: rio Negro at Anavil-hanas, 2°42' S 60°45' W. – MZUSP 29130, 2, 24.1-56.2 mmSL; Roraima: Marará: rio Branco, 1°30'N 61°16' W. –MZUSP 29144, 7, 39.6-57.2 mm SL; Amazonas: Tefé: rioTefé at Jauari-Atuuba, 3°22' S 64°43' W. – MZUSP 29150,18, 13.8-50.8 mm SL; Amazonas: Jurupari: rio Tefé. –MZUSP 29151, 5, 42.8-64.8 mm SL; Amazonas: Araná-Tuba, rio Tefé. – MZUSP 36068, 3, 162.7-216.0 mm SL;Amazonas: rio Japurá, at Lago Amanã, 2°29' S 64°37' W.– MZUSP 36073, 1, 212.7 mm SL; Amazonas: rio Japurá,2°45' S 64°30' W. – MZUSP 36075, 3, 141.4-220.6 mm SL;MZUSP 36079, 4, 182.0-236.2 mm SL; Amazonas: rio Japurá at Lago Amanã. – MZUSP 36930, 1, 221.0 mmSL; MZUSP 37439, 2, 220.0-230.5 mm SL; MZUSP 37469,1, 394.3 mm SL; Mato Grosso: Vila Bela da Santíssima

Trindade: rio Guaporé. – MZUSP 48378, 150.6-

152.0 mmSL; Pará: Oriximiná: rio Trombetas. – MZUSP 48381, 4,170.4-196.0 mm SL; Amazonas: rio Solimões at Lago


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Janauacá. – MZUSP 48384, 3, 69.6-74.2 mm SL; Ama-zonas: Lago Janauacá. – MZUSP 43444, 3, 65.2-88.4 mmSL; Amazonas: Tapera: rio Negro, 0°12' S 64°04' W. –MZUSP 48386, 1, 112.7 mm SL; Amazonas: Manaus: rioPreto da Eva. – MZUSP 48387, 1, 139.9 mm SL, Ama-zonas: Santo Antônio do Iça: rio Iça. – MZUSP 48698,

1, 140.5 mm SL; Amazonas: Fonte Boa: Igarapé Tomé,tributary of rio Solimões, 2°31' S 66°06' W. – MZUSP59059, 11, 166.2-220.6 mm SL; Amazonas: Tapera: rioNegro. – MZUSP 63209, 1, 47.0 mm SL; Amazonas: Tefé:rio Tefé, at Lago Amanã. – MZUSP 63408, 2, 255.4-267.0 mm SL; Mato Grosso: Aripuanã: rio Aripuanã,9°34'45" S 59°25'19" W. – MZUSP 83552, 14, 182.0-299.0 mm SL; Amazonas: rio Marauiá, rio Negro basin,0°24' S 65°12' W. – MZUSP 84260, 1, 158.4 mm SL;Rondônia: Montenegro: tributary of rio Jamari, rioMadeira basin. – MZUSP 91438, 2, 237.3-280.4 mm SL;Amazonas: Tapera: rio Negro. – MZUSP 92358, 1,172.2 mm SL; Amazonas: rio Tiquié, rio Negro basin,0°10'N 69°07' W. – MZUSP 101529, 1, 173.4 mm SL; Pará:Monte Dourado: rio Jari, 0°37'52" S 52°31'35" W. –MZUSP 101537, 1, unmeasured; Amapá: Laranjal do

Jari: rio Iratapuru, tributary of rio Jari. – MZUSP 101543,3, 101.8-108.9 mm SL; Amapá: Laranjal do Jari: rio Jari,0°34'16" S 52°34'44" W. – MZUSP 101696, 1, 226.4 mmSL; Pará: Monte Dourado: rio Jari, 0°35'38" S 52°38'55" W.– MZUSP 101763, 1, 227.9 mm SL; Pará: Monte Doura-do: rio Jari, 0°39'45" S 52°31'40" W. – MZUSP 101740, 1,153.8 mm SL; Pará: Monte Dourado: rio Jari, 0°37'01" S52°31'33" W. – MZUSP 101766, 3, 85.6-116.0 mm SL;Amapá: Laranjal do Jari: rio Jari, 0°33'59" S 52°34'40" W.

– MZUSP 101902, 99.5-

105.8 mm SL; Amapá: Laranjaldo Jari: rio Jari, 0°46'54" S 52°31'48" W. – MZUSP 101903,207.9 mm SL; Pará: Monte Dourado: rio Pacanari,tributary of rio Jari, 0°39'43" S 52°31'45" W. – MZUSP101904, 1, 97.1 mm SL; Pará: Monte Dourado: rio Irata-puru, tributary of rio Jari, 0°34'03" S 52°34'41" W. –MZUSP 101905, 1, 212.9 mm SL; Pará: Monte Dourado:rio Jari, 0°39'15" S 52°31'13" W. – MZUSP 102417, 1,195.7 mm SL; Amapá: Laranjal do Jari: rio Jari. – MZUSP103178, 1, 207.0 mm SL; Amazonas: rio Japurá, 1°53' S67°00' W. – MZUSP 103228, 1, 251.9 mm SL; Amazonas:Canta Galo, rio Negro. – MZUSP 103242, 4, 77.0-225.0 mm SL; Amapá: Laranjal do Jari: rio Iratapuru,tributary of rio Jari, 0°33'59" S 52°34'43" W. – MZUSP103453, 1, 232.9 mm SL; Pará, Monte Dourado: rio Jari,0°33'59" S 52°34'43" W. – MZUSP 104725, 1, 344.0 mmSL; Pará; Monte Dourado: rio Jari. – MZUSP 104744, 1,252.3 mm SL; MZUSP 104857, 1, 213.2 mm SL; Pará:Monte Dourado: rio Jari, 0°36'9" S 52°31'35" W. – MZUSP104835, 1, 170.2 mm SL; Amapá: Laranjal do Jari: Ig-arapé Arapiranga, tributary of rio Jari, 0°48'05" S52°27'20" W. – MZUSP 104888, 1, 147.7 mm SL; Amapá:Laranjal do Jari: rio Iratapuru, tributary of rio Jari, 0°33'59" S 52°34'43" W. – MZUSP 112000, 3, 84.2-103.1 mmSL; Pará: Porto de Moz: rio Xingu. – MZUSP 112211, 1,

57.5 mm SL; Amazonas: rio Negro at Ilha do Buiu-Açu.– MZUSP 112212, 1, 55.5 mm SL; Amazonas: rio Negro

below rio Daraá, 0°28' S 64°46' W. – MZUSP 112213, 4,

35.7-52.2 mm SL; Amazonas: Barcelos: rio Negro, 0°58' S62°57' W. – MZUSP 112214, 1, 136.6 mm SL; Amazonas:rio Uaupés. – MZUSP 112210, 4, 19.2-24.9 mm SL;Amazonas: Santa Isabel do Rio Negro: rio Negro, 0°31' S64°50' W. – UF 100622, 1, 180.0 mm SL; Rondônia: rio

Jamari, tributary of rio Madeira. FRENCH GUIANA:

BMNH 1909.9.4.1, 1, 158.5 mm SL; San Lorenzo: Maro-ni river. GUYANA: ANSP 176829, 2, 147.5-157.4 mm SL;Essequibo: Essequibo river at Paddle Rock Campsite.– ANSP 176830, 1, 144.8 mm SL; Essequibo: BurroBurro river at Water Dog Falls Campsite. – BMNH1911.10.3.1.475, 3, 68.9-230.0 mm SL; Bartica: Essequiboriver. – BMNH 1934.9.12.317-318, 1, 146.4 mm SL; Es-sequibo river at Bartica. – BMNH 1972.7.27.148-150, 3,116.0-246.0 mm SL; Essequibo river at Demerara. – CAS116628, 10, 55.3-171.5 mm SL; Essequibo river at Bar-tica Rock. – MZUSP 108841, 1, 208.7 mm SL; Potaro-Siparuni: Kuribrong river, 5°23'48" N 59°32'00" W. –

MZUSP 108874, 1, 175.7 mm SL; Potaro-Siparuni:Kuribrong river, 5°32'46" N 59°17'03" W. – MZUSP108860, 1, 169.3 mm SL; Potaro-Siparuni: Kuribrongriver, 5°32'04" N 59°18'37" W. PERU: ANSP 21467, holo-type of L. holostictus, 101.1 mm SL; Peruvian Amazon.– ANSP 21468, 1, 72.5 mm SL; Peruvian Amazon (pos-sibly collected with holotype of L. holostictus). – ANSP137831, 8, 153.0-182.0 mm SL; Loreto: Iquitos: río Nanay.– CAS 117234, 1, 91.6 mm SL; Loreto: río Ampiyacunear Pebas. – MZUSP 26723, 1, 38.6 mm SL; Ucayali:Colonel Portillo: río Ucayali. SURINAME: UF 16266, 1,110.4 mm SL; Marowijne: Marowijne river. – MZUSP13210, 4, 137.9-179.2 mm SL; Brokopondo: Sara creek.

– MZUSP 99416, 2, 81.5-

102.2 mm SL; Maroni: Marowi- jne river. VENEZUELA: CAS 20164, 3, 59.2-65.8 mm SL;Amazonas: río Orinoco, between Atabapo and AturesRapids. – CAS 20165, 1, 98.0 mm SL; Amazonas: ríoAtabapo, tributary of río Orinoco. – CAS 20166, 1,80.7 mm SL; río Orinoco at Chono Salata Rapids. – CAS20168, 1, 143.1 mm SL; Bolivar: río Orinoco at Caicara.– CAS 70751, 1, 245.0 mm SL; Amazonas: río Orinocoat Laja Suspiro. – FMNH 93087, 1, 157.5 mm SL; ríoCaroní, tributary of río Orinoco. – MCNG 21340, 2,120.0-161.5 mm SL; Bolivar: Departamento Sucre: ríoCaura at Jubillat, 6°38'00" N 64°37'00" W. – MCNG 51526,3, 75.1-91.6 mm SL; Apure: río Cunavichito at PuenteSan Felipe. – MCNG 40686, 1, 177.4 mm SL; Apure:Departamento Pedro Camejo: río Cinaruco, 6°32'20" N67°24'08.5" W. – MCNG 53536, 1, 137.7 mm SL; Apure:Pedro Camejo: Caño La Guardia at Puerto La LagunaMorocoto. – MCNG 52441, 2, 170.1-171.0 mm SL; Apure:Pedro Camejo: Caño La Guardia, downstream of La-guna Morocoto. L. jatuncochi: BRAZIL: INPA 10607, 1, 145.2 mm SL;Rondônia: rio Jamari, tributary of rio Madeira. – INPA10609, 1, 177.3 mm SL; Rondônia: rio Jamari. – INPA11012, 2, 196.1-202.1 mm SL; Rondônia: rio Aripuanã.– INPA 12239, 16, 22.8-78.7 mm SL; Amazonas, rio

Madeira. – MZUSP 100574, 2, 87.0-

87.3 mm SL; MatoGrosso: Aripuanã: rio Aripuanã, tributary of rio Ma-deira. – MZUSP 101034, 1, 182.6 mm SL; Mato Grosso:



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Aripuanã: rio Aripuanã, 10°10'06" S 59°26'50" W. – UF100625, 1, 140.0 mm SL; Rondônia: rio Jamari, tributaryof rio Madeira. L. pearsoni : BOLIVIA: ANSP 69069, holotype,97.2 mm SL; río Chimore at Boca Chapare. ECUADOR:FMNH 102144, 1, 157.5 mm SL; Napo: río Arajuno,

1°04'30" S 77°32'00" W. – FMNH 102145, 2, 150.0-

184.0 mm SL; Napo: río Arajuno, 1°05'24" S 77°34'18" W.PERU:ANSP 143800, 1, 160.0 mm SL; Amazonas: Madrede Dios: río Manú. – ANSP 180841, 1, 80.7 mm SL;Amazonas: Madre de Dios: río Tahuamano. – FMNH84279, 1, 122.0 mm SL; río San Alejandro. L. tigrinus: BRAZIL: MCZ 20446, 2 syntypes, 138.3-143.9 mm SL; Goiás: exact locality unknown. – CAS70447, 3, 91.3-146.0 mm SL; Goiás: Santo Antônio: rioTocantins. – INPA uncatalogued, 8, 89.0-122.5 mm SL;Pará: Altamira: rio Xingu at Cachoeira do Espelho,3°39'05" S 52°22'43" W. – MCP 19738, 1, 111.5 mm SL;Goiás: Minaçu: rio Tocantins. – MCP 33962, 3, 100.0-118.0 mm SL; Goiás: Santa Rosa Meia Ponte: rio MeiaPonte, tributary of rio Paranaíba. – MZUSP 99495, 2 CS,82.5-112.8 mm SL; Pará: Jacareacanga: rio Teles Pires,tributary of rio Tapajós, 9°20'24" S 56°46'33" W. L. yophorus: COLOMBIA: CAS 61680, holotype,163.9 mm SL; Barrigón: río Meta, Orinoco basin. – ANSP135435, 1, 222.0 mm SL; Meta: río Negrito. VENEZUELA:ANSP 160347, 1, 99.3 mm SL; Bolivar: confluence of ríoCaura and río Orinoco. – FMNH 3765, 1, 123.3 mm SL;Bolivar: Ciudad Bolivar. – FMNH 85504, 7, 50.0-61.8 mmSL; Apure: río Aruaca. – FMNH 104001, 1, 54.8 mm SL;Barinas: río Anaro, close to mouth of río Suripa. – FMNH

104004, 1, 59.0 mm SL; Barinas: Caño La Indiacita,tributary of río Suripa. – FMNH 104006, 2, 75.9-76.5 mmSL; Barinas: Caõ Sucopo, tributary of río Suripa. – MCZ59632, 1, 128.3 mm SL; Lagoon close to río Orinoco,8°41'N 62°00' W. – UF 36167, 4, 45.5-72.6 mm SL; Apure:río Orinoco basin.

Acknowledgments

For critical review of the manuscript we thank BrianSidlauskas, Fernando Jerep and an anonymous re-

viewer. For various discussions we thank Oscar Shiba-tta. We thank Alec Zeinad and Tiago Carvalho forproviding pictures of live specimens (Figs. 2e and 6b,respectively); and Barbara Lundrigan for the picture ofthe holotype of L. jatuncochi. For loan and assistancesduring collection visits we thank John Lundberg andMark Sabaj Perez (ANSP), Ralf Britz, James Maclaineand Oliver Crimmen (BMNH), Tomio Iwamoto andDave Catania (CAS), Kevin Swagel and Susan Mochel(FMNH), Lucia Rapp Py-Daniel and Renildo Oliveira(INPA), Francisco Provenzano (MBUCV), Otto Castillo(MCNG), Carlos Lucena, Margarete Lucena and Rob-erto Reis (MCP), Karsten Hartel and Andrew Williston(MCZ), Paulo Buckup and Marcelo Britto (MNRJ);Wolmar Wosiacki, Alberto Akama and André Netto-Ferreira (MPEG), and Larry Page (UF). This study was

financially supported by CNPq (JLOB, grant 47890/2013-9), and FAPESP (JLOB, grant 10/51250-9). Authorsreceived productivity research grants from CNPq (HAB)and Fundação Araucária (JLOB, grant 641/2014). Thiscontribution was also supported by the South AmericanCharaciformes Inventory project (FAPESP 2011/50282-7).

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Received 11 July 2015Revised 12 February 2016

Accepted 6 March 2016



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AQUATIC SCIENCES and FISHERIES ABSTRACTSBIOLIS - BIOLOGISCHE LITERATUR INFORMATION SENCKENBERG

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C O N T E N T S

Chen, Yan-Qiao, Chang-Lian Peng and E Zhang: Sinocyclocheilus guanyangensis, a newspecies of cavefish from the Li-Jiang basin of Guangxi, China (Teleostei: Cyprinidae) . 1

Cardoso, Yamila P., Florencia Brancolini, Ariel Paracampo, Marta Lizarralde, RaphaelCovain and Juan I. Montoya-Burgos: Hypostomus formosae, a new catfish species fromthe Paraguay River Basin with redescription of H. boulengeri (Siluriformes: Loricari-idae) .............................................................................................................................................. 9

Britski, Heraldo A. and José L. O. Birindelli: Redescription of Leporinus altipinnis, a seniorsynonym of L. falcipinnis, and comments on L. holostictus (Characiformes: Anostomi-dae) ................................................................................................................................................ 25

Britz, Ralf: Pillaiabrachia siniae, a new species of earthworm eel from northern Myanmar(Teleostei: Synbranchiformes: Chaudhuriidae) ..................................................................... 41

Decru, Eva, Emmanuel Vreven, Oumar Sadio and Jos Snoeks: Brycinus epuluensis, a newspecies from the Epulu River (Congo basin), Africa (Teleostei: Alestidae) ...................... 49

Liu, Shu-Wei, Jun-Xing Yang and Xiao-Yong Chen: Paralepidocephalus translucens, a newspecies of loach from a cave in eastern Yunnan, China (Teleostei: Cobitidae)................. 61

de Oliveira, Renildo R., Lucia Rapp Py-Daniel, Claudio H. Zawadzki and Jansen Zuanon:Two new Amazonian species of Ancistrus with vestigial adipose fin, with an appraisalon adipose fin loss in neotropical armoured catfishes (Teleostei: Loricariidae) .............. 67

Varella, Henrique R. and Ricardo Britzke: Apistogramma eleutheria and A. flavipedunculata,two new species of dwarf cichlids from the rio Curuá on Serra do Cachimbo, Brazil(Teleostei: Cichlidae) .................................................................................................................. 81


Volume 27 • Number 1 • April 2016

Cover photographPillaiabrachia siniae (photograph by Ralf Britz)

Ralf Britz(this volume pp. 41-47)

Documents

Britski Birindelli 2016 Leporinus Altipinnis