Three New Species of Fishes of the

Genus Nannostomus from the Brazilian

States of Para and Amazonas

(Teleostei: Lebiasinidae)

STANLEY H.VWEITZMAN

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 263
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 26 3

Three New Species of Fishes of the

Genus Nannostomus from the Brazilian

States of Para and Amazonas

(Teleostei: Lebiasinidae)

Stanley H. Weitzman

SMITHSONIAN INSTITUTION PRESS

City of Washington

1978
ABSTRACT

Weitzman, Stanley H. Three New Species of the Genus Nannostomus from the

Brazilian States of Para and Amazonas (Teleostei: Lebiasinidae). Smithsonian

Contributions to Zoology, number 263, 14 pages, 10 figures, 1978.—The genus

Nannostomus was recently reviewed by Weitzman and Cobb (Smithsonian

Contributions to Zoology, number 186, 1975). The three new species described

here are based on specimens found in a survey of the collections of the Museu

de Zoologia da Universidade de Sao Paulo, Brazil. The new species are N. nitidus

from Igarape Candiru-Mirim near Badajos, Rio Capim, Para; N. britskii from

an igarap6 of Lago Jose-Ac,u, Parintins, Amazonas; and AT. limatus from an

igarape tributary to Lago Mapara near Santarem, the Rio Maica near Santarem,

and the Lago Jacupa near Oriximina, all in the State of Para.

As with the other species in the genus, the phylogenetic relationships of the

new ones remain unclear. Nannostomus britskii shares a complex apomorphic

color pattern with N. eques suggesting a possible relationship between the two,

but N. britskii lacks the apomorphic caudal structure and probably the swimming position shared by N. eques and N. unifasciatus. Possible relationships

among these three species need further investigation. No close relationships

with other species of Nannostomus are suggested for N. limatus or N. nitidus.

A key is provided for the identification of all species in the genus.

RESUMO

Sao descritas tres especies novas do genero Nannostomus da parte inferior

da Bacia Amazonica, Brasil: N. nitidus, da area do Rio Capim no Estado do

Para, N. britskii do Lago Jose Acu, Parintins, Estado do Amazonas e N. limatus

dos lagos Mapara e Jacupa e Rio Maica, Estado do Para. As novas especies

elevam para 14 o numero total de especies reconhecidas, das quais 6 foram

descritas nos ultimas 25 anos em um genero conhecido a mais de 100 anos.

Isto mostra que, possivelmente, mais especies possam ser descobertas nas complexas faunas de peixes das bacias Amazonica e do Orenoco, bem como nos

rios costeiros do este situados entre elas.

As relacoes entre Ar. limatus e N. nitidus com as outras especies do genero

nao sao claras. N. britskii, entretanto, possui em comum com Ar. eques um

padrao de colorido aparentemente especializado e nao encontrado nos outros

membros do genero. Este fato parece indicar que existe uma relacao muito

maior entre estas dues especies do que com as demais especies do genero. Esta

hipotese, entretanto, precisa ser investigada, tendo em vista que Weitzman

(1966) e Weitzman e Cobb (1975) sugeriram, com base na morfologia da nadadeira caudal e na postura de natacao, que N. eques e N. unifasciatus sao mais

relacionados entre si. Nao ha evidencia de que Ar. britskii e N. unifasciatus

sejam intimamente relacionados, uma vez que N. britskii nao possui a morfologia especializada da nadadeira caudal de N. eques e de N. unifasciatus e

provavelmente tambem nao possui a posi^So caracteristica "focinho para cima"

durante a natacao, peculiar a estas duas especies. As relacoes filogeneticas da

maior parte das especies do genero permanecem obscuras.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Weitzman, Stanley H.

Three new species of fishes of the genus Nannostomus from the Brazilian states of Para and

Amazonas (Teleostei: Lebiasinidae)

(Smithsonian contributions to zoology ; 263)

Bibliography: p.

1. Nannostomas. 2. Fishes—Classification. 3. Fishes—Brazil. I. Title: Three new species of

fishes of the genus Nannostonuis .. . II. Series: Smithsonian Institution. Smithsonian contributions to zoology ; 263.

QL1.S54 no. 263 [QL638.L37] 591'.08s [59T58] 77-10751
Contents

Page

Introduction 1

Artificial Key to the Species of Nannostomus 2

Nannostonmus nitidus, new species 3

Nannostomus britskii, new species 5

Nannostomus limatus, new species .•. 9

Discussion 13

Literature Cited 14

in

Three New Species of Fishes of the

Genus Nannostomus from the Brazilian

States of Para and Amazonas

(Teleostei: Lebiasinidae)

Stanley H. Weitzman

Introduction

A recent survey of the collections of the Museu

de Zoologia da Universidade de Sao Paulo for

lebiasinid characoid fishes led to the discovery of

three new species of the genus Nannostomus Giinther, all collected as a result of the Expedicao

Permanente de Amazonia under the direction of

Dr. Paulo Vanzolini. Nannostomus, known from

Guyana, Surinam, French Guiana, the Amazon

basin in Brazil, Peru, Bolivia, Venezuela, and Colombia, and part of the Orinoco basin in Venezuela, was last reviewed by Weitzman and Cobb

(1975). They recognized 11 species, of which three

had been described as new since 1953. Thus, counting the three species described here, three-sevenths

(about 43%) of the recognized species of Nannostomus have been described within the last 25 years

in a genus that has been known for a little over

100 years. Considering the low geographical density

and irregularity of fish population samples in the

varied and extremely extensive forested areas of

the Amazon and Orinoco basins to which Nannostomus appears confined, it seems possible that several more species may remain to be discovered.

Several people have made the study of these

Stanley H. Weitzman, Department of Vertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

specimens possible. Dr. Paulo Vanzolini, director

of the Museu de Zoologia da Universidade de Sao

Paulo, graciously allowed me to examine and study

characoids from the collections in the Museu de

Zoologia and to sort through the collections made

by the Expedicao Permanente da Amazonia

(E.P:A.), which is supported by the Fundacao de

Amparo a Pesquisa do Estado de Sao Paulo. Drs.

Heraldo Britski and Naercio A. Menezes of the

Museu de Zoologia da Universidade de Sao Paulo

provided much assistance and counsel during my

examination of the collections in their care. Furthur assistance in locating, sorting, and examining

specimens in the collections of the Museu de Zoologia was provided by Marilyn Weitzman, William

L. Fink, and Sara V. Fink. Funds for examination

of the collections at the Museu de Zoologia were

provided by the Smithsonian Amazon Ecosystem

Research Program under the direction of Dr. Clifford Evans, Smithsonian Institution. Naercio Menezes provided the Portuguese translation of the

"Resumo." Drs. Robert H. Gibbs, Jr., and William

L. Fink read the manuscript and offered valuable

suggestions. I mucli appreciate the support of all

these people and associated institutions.

Collections are deposited in the following museums: Academy of Natural Sciences of Philadelphia (AXSP), British Museum (Natural History)

(BMNH), Museu de Zoologia da Universidade de

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sao Paulo (MZUSP), and National Museum of

Natural History (USNM, formerly United States

National Museum).

The methods of making counts and measurements and the color pattern terminology below are

the same as those described by Weitzman (1966)

and Weitzman and Cobb (1975). Measurements are

given here as thousandths of standard length. In

Weitzman (1966) the arithmetic mean was given,

followed in parentheses by the range. Herein the

figure for the holotype is given first in brackets when

different from other specimens and is followed by

the range and the mean of the paratypes. The Weberian apparatus and the terminal half centrum are

included in the vertebral counts.

In order to bring the key of Weitzman and Cobb

(1975) up to date, it is modified here to include the

three new species described below.

Artificial Key to the Species of Nannostomus

1. Principal caudal-fin rays 10/9, their distribution in dorsal and ventral caudal lobes as in

most characoids, 10 ending in dorsal lobe, 9 ending in ventral caudal-fin lobe 2

Principal caudal-fin rays 10/9, but 8 rays end in dorsal lobe and 11 in ventral caudal-fin

lobe 13

2. Primary horizontal stripe very indistinct. Secondary horizontal stripe very weakly developed and tertiary horizontal stripe absent. Five dark permanent blotches along sides

(blotches not fading during exposure of live fish to daylight) Nannostotnus espei

Primary horizontal stripe well developed. Secondary and/or tertiary horizontal stripes

present in some species, absent in others. One or two pale vertical or oblique bands

on sides (in life these bands ordinarily prominent and darker when fish kept in dark

or at night; oblique bands sometimes intensified during breeding behavior, especially

in females). Permanent blotches absent 3

3. Secondary horizontal stripe usually present, sometimes poorly developed or absent. Tertiary

horizontal stripe absent or poorly developed, represented by only a few black spots

associated with pelvic-fin base 4

Secondary and tertiary horizontal stripes well developed; tertiary stripes sometimes reduced

to a few dark spots anterior to pelvic-fin bases but always well developed posterior

to these fin bases 8

4. Adipose fin always present 5

Adipose fin always absent 7

5. Sensory canal present in second infraorbital bone. Snout in eye 0.9 to 1.0. Secondary dark

horizontal stripe usually poorly developed. Total vertebrae 38 or 39

Nannostotnus harrisoni

Sensory canal absent in second infraorbital bone. Snout in eye 1.0 to 1.3. Secondary dark

horizontal stripe well developed. Total vertebrae 34 to 35 6

6. Gill rakers 10 or 11 + 16 to 18. Anal fin of males never elongate, nearly reaching ventral

base of caudal fin. Anal-fin rays moderately modified, somewhat enlarged and thickened..

Nannostomus nitidus

Gill rakers 7 + 14. Anal fin of males over 24 to 25% of standard length, nearly or reaching

ventral caudal-fin base and fin rays greatly modified, flattened and expanded saggital

plane Nannostomus digrammus

7. Secondary horizontal stripe poorly developed or absent. Gill rakers 9 or 10 + 17 or 18 ..

Nannostomus beckfordi

Secondary horizontal stripe well developed. Gill rakers 8 + 14 Nannostomus bifasciatus

8. Anal fin of males long, reaching or nearly reaching ventral origin of caudal fin when

adpressed to body Nannostomus minimus

Anal fin of males short, not reaching to or nearly to ventral origin of caudal fin when

adpressed to body 9

9. Two dark horizontal stripes on back dorsal to secondary horizontal stripe, one median

extending through predorsal scales, other extending along center of second horizontal

scale row. Primary horizontal stripe divided into two distinct stripes, one concentrated

along the fourth horizontal scale row, the next along the fifth horizontal scale row.

Anal fin of males with moderately modified and thickened fin rays

Nannostomus britskii
NUMBER 263

Dark horizontal stripes absent on back dorsal to secondary horizontal stripe. Primary

horizontal stripe of one solid unit confined mostly to ventral portion of third and dorsal

of fourth horizontal scale row or to fourth scale row and dorsal portion of fifth scale

row. Anal fin of males moderately modified or apparently not modified at all (N. marginatus and N. trifasciatus) 10

10. Gill rakers 11 or 12 + 17 or 18. Simple conic teeth of inner dentary row absent

Nannostomus litnatus

Gill rakers 7 to 9 + 12 to 14. Simple conic teeth of inner dentary row 6 to 11 11

11. Scales in a lateral series 21 to 23. Adipose fin always absent Nannostotnus marginatus

Scales in a lateral series 24 to 27. Adipose fin present (sometimes absent in N. trifasciatus)

12

12. Oblique (vertical) bands broad; anterior band extending posteriorly well past a vertical

line from anterior base of dorsal fin, frequently past a vertical from midbase of dorsal

fin. Posterior oblique band extending posteriorly well beyond posterior base of anal fin.

Eye diameter about 1.0 in interorbital width Natmostomus trafasciatus

Oblique bands narrow; anterior band reaching but not extending past a vertical line

from anterior base of dorsal fin. Posterior oblique band not reaching a vertical from

posterior base of anal fin. Eye diameter about 0.8 in interorbital width

Nanostomus marilynae

13. Scales 28 to 30 in lateral series. Perforated scales in lateral line 2 to 5. Adipose fin present.

Gill rakers 9 + 14. Inner tooth row of dentary absent; teeth in outer dentary row 6 to 7..

Nannostomus unifasciatus

Scales 24 or 25 in lateral series. No perforated lateral-line scales. Adipose fin present or

absent. Gill rakers 16 + 24. Teeth in inner dentary row 12; teeth in outer dentary row 9

Nannostomus eques

Nannostomus nitidus, new species

FIGURES 1, 2

HOLOTYPE.—MZUSP 12920, male, SL 34.7 mm;

Brazil, State of Para, Igarape Candiru-Mirim near

Badaj6s, Rio Capim, 29 Aug 1970, Naercio Menezes, E.P.A.

PARATYPES.—18, MZUSP 12921-12938, SL 16.3

to 33.3 mm; same data as holotype. 8, USNM

216204, SL 17.0 to 34.5 mm; same data as holotype.

5, ANSP 137388, SL 18.3 to 34.1 mm; same data

as holotype. BMNH 1976-11-8-1-5, SL 18.5 to

35.2 mm; same data as holotype.

DIAGNOSIS.—Primary and secondary horizontal

stripes present, all others absent. Nocturnal oblique

bands present, narrow, anterior band not extending

posteriorly much beyond a line extending vertically

through anterior dorsal-fin origin; permanent

blotches absent. Adipose fin present. Scales in lateral series 23 to 25, usually 24. Perforated lateralline scales 0 to 2. Total vertebrae 34 or 35. Gill

rakers 10 to 11 + 16 to 18. Simple conic teeth in

inner dentary row 2 or 3, laterally placed. Principal

caudal-fin rays 10/9, 10 terminating posteriorly in

dorsal lobe, 9 in ventral lobe. Eye about 100 to

110% of least bony interorbital width. Body depth

FIGURE 1.—Nannostomus nitidus, M7.USP 12920, male, 34.7 mm SL, holotype,

Igarape Candiru-Mirim near Badajos, Rio Capim, Para, Brazil.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Nannostomus nitidus, USNM 216204, female, 30.5 mm SL, paratype,

Igarape Candiru-Mirim near Badaj6s, Rio Capim, Para, Brazil.

about 23 to 25% of standard length, and least

depth of caudal peduncle about 9% of standard

length. Anal fin of male moderately modified, having slightly thicker rays than that of female.

DESCRIPTION.—Body elongate, compressed in region of caudal peduncle, greatest body depth at

origin of dorsal fin [239] 235-253, x = 243; least

depth of caudal peduncle [089] 088-094, x = 090;

snout tip to origin of dorsal fin [513] 507-536, x =

519; snout tip to origin of anal fin [764] 734-782,

x = 760.

Head conic, snout obtuse. Head [242] 238-265,

x = 251; eye [079] 079-088, x = 084; least width of

bony interorbital [078] 075-079, x = 077.

Premaxillary with 6 teeth, first (anterior) 5 or 6

with 6 subequal cusps, and sixth tooth sometimes

small but with 6 cusps, median cusp largest. Maxillary toothless. Dentary with 6 teeth in outer row,

each bearing 6 subequal cusps. Inner dentary row

with 2 or 3 conic teeth, placed laterally in jaw

(two alizarin preparations).

Gill rakers [11 + 18], 10 + 16 in one and 10 +

17 in a second alizarin preparation.

Dorsal fin ii, 8; anal fin iii, 8; pectoral fin i, 11;

pelvic fin ii, 7; and principal caudal-fin rays 10/9

posteriorly terminating in dorsal lobe and 9 in

ventral lobe in all specimens. Adipose fin present

in all specimens.

Scales belonging to third horizontal series and

lying ventral to dorsal fin having anterior field

with 3 radial grooves and posterior field with 2

radial grooves. Dorsal and ventral fields each with

1 radial groove. Scales in a lateral series [25] 23

to 25, x = 24.2. Lateral-line scales [0] 0 to 2, usually

on third and fourth scale horizontal scale row; 10

predorsal scales in all specimens.

Total vertebrae [35] 9 paratypes with 34 and

11 with 35.

COLOR IN ALCOHOL.—Dorsal surface of head

dark brown, more or less evenly covered with small,

dark brown chromatophores. Back dorsal to secondary horizontal stripe dark brown but not as

dark as top of head, rather densely covered with

moderate-sized brown chromatophores except for

distal free margins of scales, where chromatophores

less dense than central portions of scales. Each

scale bordered by a pale brown band. No dark

brown longitudinal stripes occur dorsal to secondary stripe. Primary horizontal stripe very dark

brown, nearly black, consisting of a dense aggregation of large, very dark brown chromatophores.

Primary stripe originating anteriorly at tips of

upper and lower jaws, extending across orbital

bones to reach anterior border of eye, continuing

across eye, postorbital and preopercular areas and

operculum to include a few dark brown chromatophores scattered on fleshy opercular flap. On body,

primary stripe extending from under fleshy opercular flap, beginning at cleithrum and supracleithrum, and continuing posteriorly over ventral half

of third horizontal scale row and dorsal half of

fourth horizontal scale row, ending on scales

covering base of ventral caudal-fin lobe. Primary

stripe apparently not extending onto fin rays of

ventral caudal-fin lobe. Secondary dark horizontal

stripe extending from just dorsal to opercular bone

(where it joins head dorsally) to across middle of

second horizontal scale row. Secondary horizontal

stripe fading to color of back just posterior to a
NUMBER 263

vertical line extending ventrally from posterior

base of dorsal fin. Secondary stripe in preserved

specimens consisting of a cluster of very dark brown

chromatophores concentrated at center of each

scale of second horizontal scale row, forming a

series of spots. Pale yellow to white stripe, situated

between primary and secondary stripes, occupying

dorsal half of third scale row and ventral half of

second scale row, and containing a few brown

chromatophores scattered along scale margins. Pale

stripe also contains dorsal portions of anterior and

posterior oblique bands. Anterior oblique band in

area of pale stripe extending ventroanteriorly from

mostly seventh, eighth, and ninth scales of third

horizontal scale series ventral to primary stripe,

where it covers all of ventral portions of fifth and

sixth scales (counting dorsal postcleithrum as a

scale) of fourth longitudinal scale row. Posterior

oblique band covering about 17th and 18th scales

of third horizontal scale row, and extending ventrally to cover parts of 15th through 18th scales

of fourth longitudinal scale row, which extends

just ventral to primary horizontal stripe. Sides ventral to primary horizontal stripe nearly devoid of

pigment. No evidence of a tertiary horizontal stripe

and only a very few scattered, small, dark brown

chromatophores around vent in either sex. All

rayed fins except caudal fin with small brown chromatophores along edges of bony rays producing

hyaline but slightly dusky fins. Pectoral, and especially pelvic and anal, fins with more distal portions of ray, especially ray tip, with dark brown

chromatophores, forming faintly darkened edges

to fins. Caudal fin nearly hyaline except for narrow black edge above dorsal and ventral borders.

Life colors unknown.

ETYMOLOGY.—The name nitidus is a Latin adjective for "neat, elegant, or shining."

RELATIONSHIPS.—Although Nannostomus nitidus

keys to Nannostomus digrammus in the artificial

key of Weitzman and Cobb (1975), no relationship

between N. nitidus and N. digrammus is hypothesized, especially since at large sizes (to 34 and

nearly 35 mm SL), males of N. nitidus lack the

highly specialized fin rays and increased length ot

the anal fin found in males of N. digrammus over

24 to 25 mm in standard length. Nannostomus

nitidus further differs from N. digrammus in having 10 to 11 + 16 to 18 (rather than 7 + 14) gill

rakers. The two species differ in the details of

color pattern; for example, N. digrammus has the

primary horizontal stripe continued as a wedge

onto the caudal fin, whereas the wedge is absent

on this fin in N. nitidus. I found nothing in the

color pattern or morphometric and meristic characters that suggests any hypothesis of sister group

relationship on the part of nitidus with any other

species of Nannostomus.

Nannostomus britskii, new species

FIGURES 3, 4

HOLOTYPE.—MZUSP 7795, male, SL 22.8 mm;

Brazil, State of Amazonas, an igarape" of Lago

Jos£-Ac,u, Parintins, Amazon River, 11-12 Dec

1967, Heraldo Britskii, E.P.A.

PARATYPES— MZUSP 13072, one, female, SL 24.1

mm; same data as holotype. USNM 216203, one,

female, SL 23.0 mm; same data as holotype.

DIAGNOSIS.—Primary, secondary, and tertiary

horizontal stripes well developed, with an additional stripe (quaternary) on back dorsal to secondary stripe. Nocturnal oblique bands present,

anterior band narrow, not extending much behind

anterior origin of dorsal fin; permanent blotches

absent. Adipose fin absent. Scales in a lateral series

21 or 22. Simple conic teeth in inner dentary row

12, teeth in outer dentary row 5. Principal caudalfin rays 10/9, those ending in dorsal lobe 10, those

in ventral lobe 9. Eye about 70 to 80% of least

bony interorbital width. Body depth about 22 to

25% and caudal peduncle depth about 11 to 12%

of standard length. Anal fin of male moderately

modified with fin rays somewhat thicker than

those of female.

DESCRIPTION.—Body elongate, robust; compressed

in region of caudal peduncle. Greatest body depth

at origin of dorsal fin [241] 226-241, x = 237;

least depth of caudal peduncle [114] 114-116,

x = 114; snout tip to origin of dorsal fin [553]

548-577, x = 559; snout tip to origin of anal fin

[768] 765-788, x = 774.

Head conic, snout obtuse in horizontal and vertical profile. Head [280] 265-280, x = 273; eye [101]

100-101, x = 100; least width of bony interorbital

[105] 104-109, x = 106.

Premaxillary with 6 teeth, all with 4 subequal

cusps except first (anterior) with 3 subequal cusps.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 5.—Nannostomus britskii, MZUSP 7795, male, 22.8 mm SL, holotype,

igarape of Lago Jose-A^u, Parintins, Amazon River, Amazonas, Brazil.

Maxillary with 1 tooth bearing 3 cusps, the middle

cusp the longest. Dentary with 5 teeth in outer row,

each tooth bearing 4 or 5 more or less subequal

cusps. Inner row of dentary teeth simple, conic

with 12 teeth placed along entire inner margin

of jaw.

Gill rakers 17 + 23.

Dorsal fin ii, 8; anal fin iii, 8; pectoral fin i, 9;

pelvic fin ii, 7; principal caudal-fin rays 10/9 with

10 ending in dorsal lobe and 9 terminating in ventral lobe in all specimens. Adipose fin absent in

all specimens.

Scales of third horizontal scale series ventral to

dorsal fin having anterior field with 3 radial

grooves, posterior field with 2 radial grooves; dorsal

and ventral fields each with 1 radial groove. Scales

in lateral series [22] 21 and 22; [0] 0 or 1 perforated

lateral-line scales; 10 predorsal scales in all specimens.

Total number of vertebrae 32 in all specimens.

COLOR IN ALCOHOL.—Dorsal surface of head medium brown, covered with fairly dense, moderatesized brown chromatophores except along midline

of braincase, which is pale brown. Back dorsal to

secondary horizontal stripe pale brown with denser

aggregation of dark brown chromatophores through

center of predorsal scales and first lateral scale row,

forming respectively a median predorsal stripe and

a fourth horizontal stripe very similar to that ot

Nannostomus eques. Median dorsal stripe and

fourth horizontal stripe usually difficult to distinguish posterior to dorsal fin. Primary horizontal

stripe dark brown and extending from snout tip

and lower jaw posteriorly across first (anterior) orbital bone, through eye, across preoperculum and

operculum and terminating at posterior border of

bony operculum. Primary stripe not extending onto

fleshy opercular flap. On body, primary stripe beginning under fleshy opercular flap at articulation

of supracleithrum and cleithrum and extending

posteriorly across fourth horizontal scale row to

terminate at base of caudal fin. As in N. eques

(Figure 5) fifth horizontal scale row also strongly

pigmented with dark brown chromatophores, forming in N. britskii a second, and separate, ventral

portion of primary horizontal stripe. This ventral

portion beginning from just anterior to dorsal base

of pectoral fin or at ventral border of opercular

bone and extending posteriorly across fifth horizontal scale row to base of caudal fin. Both fourth

and fifth horizontal scale rows with dark brown

chromatophores concentrated in centers of scales.

In N. eques pattern of primary stripe often like

that of N. britskii in preserved specimens but in

living (and some preserved specimens) pigment of

these scale rows may merge to form one horizontal

stripe, especially posteriorly on caudal peduncle.

Brown chromatophores extending from these scale

rows onto proximal one-third of ventral lobe of

caudal fin of N. britskii, not onto nearly entire fin

as in N. eques. Secondary horizontal stripe extending from dorsal border of operculum posteriorly

through ventral half of second horizontal scale row

and dorsal part of third horizontal scale row to

caudal-fin base. Margins of scales in area of secondary stripe dark brown. Secondary stripe very pale
NUMBER 263

posterior to posterior oblique band. Pale stripe,

occupying central area of third horizontal scale

row and between primary and secondary stripes,

with very few brown chromatophores. Almost no

dark brown chromatophores in center of scales of

third horizontal scale row except in areas of oblique

bands that occur on eighth and ninth scales of

third horizontal scale row and 16th, 17th, and part

of 18th scales of third horizontal scale row. Anterior oblique band extending anteroventrally to

reach ventral to both rows of dark scales of primary

horizontal stripe, darkening area of scales of sixth

and seventh horizontal scale rows. Posterior oblique

band not extending ventrally below second row of

dark scales of primary horizontal stripe. Sides ventral to primary horizontal dark stripe pale brown,

without dark brown chromatophores except in areas

of anterior oblique band and tertiary horizontal

stripe. Tertiary stripe extending as dark brown pigment areas in centers of scales of sixth horizontal

scale row from posterior termination of operculum

or just posterior to ventral origin of pectoral and

to just dorsal to base of pelvic fin to immediately

anterior to anus.

Dorsal and pectoral fins with a few small, dark

brown chromatophores scattered along borders of

fin rays. Pelvic fin with same distribution of dark

brown chromatophores but with a more concentrated collection of somewhat larger dark brown

chromatophores along third through fifth or sixth

branched rays. Anterior first and second unbranched

anal-fin rays with large, dark brown chromatophores. Distal portion of third unbranched ray,

first branched ray, and base of second branched ray

hyaline, without dark brown chromatophores. Remainder of anal-fin rays with large dark brown

chromatophores, and basal one-half of fifth through

eighth ray nearly black. Dorsal lobe of caudal fin

hyaline (partly missing in male), except at base

with scattered large, dark brown chromatophores.

Life colors unknown.

ETYMOLOGY.—The name britskii is for Dr. Heraldo Britski, Curator of Fishes at the Museu de

Zoologia da Universidade de Sao Paulo, Brazil, and

collector of the species of Nannostomus herein described as new.

RELATIONSHIP.—The possible phylogenetic relationships of this species are not clear. In the artificial key in Weitzman and Cobb (1975) it keys to

Nannostomus marginatus, having all 10 principal

caudal-fin rays of the dorsal lobe ending in that

lobe, having no permanent blotches, having three

or more well-developed horizontal stripes, having a

relatively short anal fin in the male, and having

21 to 22 scales in a lateral series. It has a color

pattern strikingly different, however, from that of

N. marginatus (compare Figures 2 and 3 with those

of N. marginatus in Weitzman and Cobb (1975,

figs. 22, 23, and 24), and remarkably like that of

N. eques (Figure 5). Also, in its gill-raker count

(17 + 23), N. britskii is close to that (16 + 24) of

N. eques (see also below). Nannostomus marginatus

has a gill-raker count of 7 + 12. Nannostomus britskii further differs from N. marginatus in having

5 dentary teeth in the outer row (versus 7 to 8 in

N. marginatus), 12 teeth in the inner dentary row

(versus 6 or 7 in N. marginatus), none or 1 perforated lateral-line scales (versus 3 to 5, usually 4,

FIGURE 4.—Nannostomus britskii, MZUSP 13072, female, 24.1 mm SL, paratype,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 5.—Nannostomus eques Steindachner, USNM 216552, male, 32-4 mm SL, an

igarape of the varzea at Tapurucuara, Rio Negro, Amazonas, Brazil.

in N. marginatus), and a moderately modified anal

fin in the male (there being little or no observable

morophological sexual dimorphism in the anal fin

of N. marginatus, although there is functional dimorphism in which the male's anal fin is "cupped"

around the vent of the female during spawning).

Nannostomus britskii and N. eques share a complicated and apparently specialized color pattern

(in preservative and presumably in life; see color

description above and compare Figures 3, 4, and

5). They also share a high and presumably derived

total gill raker count (a total of 40 on the first

arch). See Weitzman (1966) for counts and measurements of N. eques. The highest gill-raker count

found in other species of Nannostomus is 27 to 30

(N. espei and N. beckfordi). Nannostomus britskii

also shares with N. eques a few reductive characters

absent or not found so greatly reduced in some

other species of Nannostomus. For example, they

have none or only one perforated lateral-line scale

(2V. minimus and N. espei sometimes have only one

perforated lateral-line scale and N. digrammus has

no perforated lateral-line scales), and they share a

loss of or tendency to lose the adipose fin (but it

is also absent in N. beckfordi, N. marginatus, N.

minimus, N. bifasciatus and tends to be reduced

or occasionally lost in N. trifasciatus). These reductive characters are too likely to be convergent for

use in evaluating hypotheses of phylogenetic relationship. The gill-raker count also could be convergent although it is perhaps worth noting that

only two species, N. eques and N. britskii, have

such a high count. Evidence for a relationship between N. britskii and N. eques must at present rest

mainly on the shared (nearly identical), complicated, and apparently specialized color pattern. The

possible relationships of N. eques are complicated,

however, (see below), and the homology of the

shared color pattern is difficult to assess.

Weitzman (1966:41) and Weitzman and Cobb

(1975:4) proposed a phylogenetic relationship between N. eques and N. unifasciatus based on the

similarity of their synapomorphic caudal-fin structure and "snout-upward" (oblique) swimming position. The ventral lobe of the caudal fin of each

of these species is larger than the dorsal lobe and

contains the termination of two fin rays originating

in the dorsal lobe. These characters are apparently

associated with the specialized swimming position.

Nannostomus britskii presumably does not swim

at an oblique angle since it lacks the specialized

caudal find morphology; however, the swimming

position of N. britskii has not been recorded.

Nannostomus britskii also differs from N. eques

and N. unifasciatus in having a relatively short

snout (about 70 to 90% of the interorbital width,

112 to 125% in the other two species), by lacking

a canal in the second circumorbital (present in the

other two, in N. harrisoni, and in large specimens

of N. bifasciatus), and by having a short body (relatively long in N. eques and N. unifasciatus). The

total vertebral count for N. britskii is 32 for all

specimens. In a series of 140 specimens of N. eques

from several localities in the Amazon basin (specimens cited in Weitzman, 1966, plus 29 specimens

from Ilha Marchanteria, Rio Solimoes, south of
NUMBER 263

Manaus, Amazonas, USNM 216548, and 19 specimens from Rio Inhangapi at Inhangapi, Para,

USNM 216547, both in Brazil), the vertebral count

range was from 32 to 35 with a mean of 33.7. Only

two specimens had a count of 32 and these were

(1) from an igarape" near Sanabani, Municipio

Silves, MZUSP 7474, not far from the type-locality

of N. britskii and (2) from Lago Jacupa near Oriximina near Obidos, MZUSP 8204, over 300 km east

of the Sanabani locality. In the 19 specimens from

Sanabani the vertebral count range was from 32

to 34 with a mean of 33.1, and in the collection

from Oriximina the range was from 32 to 34 with

a mean of 33.3. There might be a shift in vertebral

numbers in the population of N. eques from Tapurucuara on the Rio Negro for in 2 samples for a

total of 16 specimens (USNM 216551 and USNM

216552) the range was from 34 to 35 vertebrae with

a mean of 34.8. Nothing about relationships can

be discovered by comparison of vertebral numbers

in these two species and it would seem that vertebral numbers in species of Nannostomus may have

typological use in distinguishing geographically

separate populations of a species and in separating

certain species from others but of little use in understanding the phylogenetic relationships of these

fishes. Many of the characters discussed above may

be easily subject to convergence and divergence and

it is difficult to assign phylogenetic significance to

any of them at present. Therefore I am unable to

propose reasonable hypotheses about the phylogenetic relationships of the three species, N. briskii, N. eques, and N. unifasciatus.

Nannostomus limatus, new species

FIGURES 6, 7, 8

HOLOTYPE.—MZUSP 8486, male, SL 29.9 mm;

Brazil, State of Para, an igarape tributary to Lago

Mapara, near Santar&n, Amazon River, 25 Dec

1967, Heraldo Britski, E.P.A.

PARATYPES.—14, MZUSP 12951-12963, SL 19.6

to 27.5 mm; same data as holotype. 7, USNM

216201 (formerly MZUSP 12964-12971), SL 19.6 to

34.2 mm; same data as holotype. 2, ANSP 137389

(formerly MZUSP 12972-12973), SL 24.9 to 27.8

mm; same data as holotype. 2, BMNH 1976—11—

8-6-7 (formerly MZUSP 12974-12975), SL 23.9 to

27.8 mm; same data as holotype. 1, MZUSP 8203,

SL 36.3 mm; Brazil, State of Para, Lago Jacupa,

Oriximina, 17 Dec 1967, Heraldo Britski, E.P.A. 1,

USNM 216200, SL 33.6 mm; same data as MZUSP

8203. 1, MZUSP 9192, SL 28.1 mm; Brazil, State

of Para, Rio Maica, Santarem, 19-27 Oct 1971,

Yasunobu Matsurura, E.P.A. 7, MZUSP 12939-

12945, SL 17.4 to 24.9 mm; same data as MZUSP

9192. 5, USNM 216202 (formerly MZUSP 12946-

12950), SL 15.5 to 27.7 mm; same data as MZUSP

9192.

DIAGNOSIS.—Primary, secondary, and tertiary horizontal stripes well developed. No additional stripes

on back. Nocturnal oblique bands present; narrow,

anterior band reaching to but not past an imaginary vertical line extending ventrally from anterior

base of dorsal fin; permanent blotches absent. Adipose fin absent. Scales in a lateral series 24 or 25.

Perforated lateral-line scales 0, 1, 2, or 3. Anal fin

FIGURE 6.—Nannostomus limatus, MZUSP 8486, male, 29.9 mm SL, holotype, an

igarape tributary to Lago Mapara, near Santarem, Para, Brazil.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 7.—Nannostomus limatus, USNM 216201, female, 31.6 mm SL, paratype, an

igarape tributary to Lago Mapara, near Santarem, Para, Brazil.

of males not elongate, moderately modified, rays

slightly thickened and margin rounded. Gill rakers

11 to 12+17 to 18. Simple conic teeth in inner

dentary row absent; 6 teeth in outer dentary row.

Principal caudal-fin rays 10/9, those posteriorly

terminating in dorsal lobe 10, those in ventral

lobe 9. Eye about 75% to 90% of interorbital

width. Body depth about 19 to 22% and caudalpeduncle depth about 9% to 10% of SL.

DESCRIPTION.—Body elongate, compressed in region of caudal peduncle. Greatest body depth at

origin of dorsal fin [217] 193-224, x = 210; least

depth of caudal peduncle [094] 087-097, x = 092;

snout tip to origin of dorsal fin [542] 498-545, x =

531; snout tip to origin of anal fin [779] 774-795,

x = 765.

Head conic, snout obtuse. Head [254] 241-271,

x = 254; eye [084] 077-091, x = 083; least width of

bony interorbital [080] 077-090, x = 083.

Premaxillary with 6 teeth, each with 5 to 6 and

occasionally 7, subequal cusps. Maxillary with one

small unicuspid or tricuspid tooth. Dentary with

6 teeth in outer row, each bearing 6 subequal cusps

(one tooth in one specimen with 5 cusps). Inner

tooth row of dentary absent (in 2 alizarin preparations).

Gill rakers 11 or 12 + 17 or 18.

Dorsal fin ii, 8 and anal fin iii, 8 in all specimens; pectoral fin [i, 10] i, 10 or i, 11 (usually i, 11);

pelvic fin ii, 7 and principal caudal-fin rays 10/9

with 10 terminating posteriorly in dorsal lobe and

9 in ventral lobe in all specimens. Adipose fin absent in all specimens.

Scales belonging to third horizontal series and

lying ventral to dorsal fin, having anterior field

with usually 3 radial grooves, posterior field usually

with 2 radial grooves (sometimes 1 or 3). Dorsal

and ventral fields each with 1 radial groove. Scales

FIGURE 8.—Nannostomus limatus, USNM 216201, juvenile, 20.1 mm SL, paratype,

an igarape tributary to Lago Mapara, near Santarem, Para, Brazil.
NUMBER 263 11

in a lateral series [25] 24 or 25; lateral-line scales

[3] 0 to 3, usually on scales 3, 4, or 5 of third scale

row; [11] 10 to 11 predorsal scales.

Total number of vertebrae [35], 17 paratypes

with 34 and 17 with 35.

COLOR IN ALCOHOL.—Dorsal surface of head medium to dark brown, more or less evenly covered

with small, brown chromatophores. Back dorsal to

secondary horizontal stripe medium brown with

rather evenly distributed moderate-sized chromatophores distributed in broad central area of scales.

Larger, slightly darker, but less densely distributed

brown chromatophores bordering posterior edge of

each scale in a broad band. No dark brown stripes

occurring dorsal to secondary horizontal stripe. Primary horizontal stripe dark brown, consisting of

densely aggregated, fairly large brown chromatophores. Primary stripe beginning anteriorly at

lower and upper jaw tips, extending across antorbital bone and dorsal anterior portion of first

infraorbital bone to reach anterior border of eye,

continuing across eye but apparently absent posterior to eye in area of circumorbital bones, continuing across operculum as a diffuse stripe, ending

on soft opercular flap. On body, primary stripe

extending from under fleshy opercular flap, beginning at articulation of deithrum and supracleithrum and continuing posteriorly across ventral

half of third horizontal scale row and dorsal halt

of fourth horizontal scale row, ending on base of

ventral lobe of caudal fin. Dorsal to primary stripe,

secondary dark brown stripe extending from just

dorsal to opercular bone along middle of second

horizontal scale row as a somewhat dense aggregation of dark brown chromatophores and ending

at anterior termination of dorsal lobe of caudal fin

at dorsal surface of caudal peduncle. Pale brown

stripe situated between primary and secondary

stripes, occupying dorsal half of third horizontal

scale row and ventral one-quarter of second horizontal scale row; this stripe with scattered brown

chromatophores except in areas of oblique bands,

where dark chromatophores are larger and denser.

Oblique bands not clearly defined in specimens at

hand, but anterior band present in seventh, eighth,

and ninth scale rows of third horizontal series and

third, fourth, and fifth scale rows of fifth horizontal

scale series (ventral to primary horizontal stripe).

Posterior oblique band occurring at 17th, 18th, and

19th scales of third horizontal scale row. Ventral

to primary horizontal stripe, posterior oblique band

extending to area of anus and across root of anal

fin. Sides ventral to primary horizontal stripe pale

brown, nearly without chromatophores, except in

areas of oblique bands. Tertiary stripe present, extending as small scattered brown chromatophores

from area of anus anteriorly to just dorsal to pelvicfin base and to near origin of pectoral fin. All rayed

fins, except caudal fin, with small brown chromatophores along borders of bony rays. Otherwise pectoral fin hyaline; second ray of dorsal fin with a

dark anterior border; pelvic fin with more brown

chromatophores at distal portion of rays than anteriorly. Second and third rays of anal fin with a

dark border continued onto distal tips of fourth,

fifth, and sixth rays; distal portion of remaining

anal-fin rays nearly hyaline. Caudal fin nearly hyaline except for dark border on dorsalmost and ventralmost principal rays. Primary stripe tending to

extend onto dorsal rays of ventral caudal-fin lobe

and may continue to their posterior termination.

ETYMOLOGY.—The name limatus is a Latin adjective for "elegant, refined, or polished."

RELATIONSHIPS.—As with most species of Nannostomus, it has proved impossible to propose reasonably testable hypotheses of the phylogenetic relationships of N. limatus to other species of Nannostomus. In the artificial key of Weitzman and Cobb

(1975), N. limatus keys to couplet 8, having an unmodified caudal fin, a relatively short anal fin in

the males, and primary, secondary, and tertiary

longitudinal stripes. Nannostomus limatus is easy

to distinguish from the three species, N. marginatus, N. trifasciatus, and N. marilynae, included

within this couplet. First, it always lacks an adipose

fin (always present in N. marilynae and nearly always present in N. trifasciatus but absent in N.

marginatus). Nannostomus limatus (Figures 7, 8,

and 9) has narrow oblique bands similar to those

of N. marilynae (Weitzman and Cobb, 1975, figs.

28 and 30) and unlike the broad oblique bands of

N. marginatus (Weitzman and Cobb, 1975, fig. 22)

and N. trifasciatus (Weitzman and Cobb, 1975, figs.

25 and 26). Also as in N. marilynae, N. limatus has

a moderately modified anal fin in the males, very

unlike the nearly unmodified anal fin of N. marginatus and N. trifasciatus. Nannostomus limatus

differs from all three species in having no posterior
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

row of conical teeth in the dentary, whereas N.

marginatus has 6 or 7 teeth, N. trifasciatus about

11, and N. marilynae about 1 to 4. Nannostomus

limatus also has a greater number of gill rakers (11

to 12 on the upper limb + 17 to 18 on the lower

limb) of the first gill arch; N. marilynae has 8 to 9

•.+ 13 to 14, N. trifasciatus his 9 + 13, and N. marginatus has 7 + 12. In color pattern, N. limatus

more closely approaches N. marilynae than the

other two species but appears to be a larger fish,

reaching at least 36.3 mm. Of over 300 specimens

of N. marilynae from various localities in the Amazon and Orinoco basins, I have found none over 23

mm in SL. Very old specimens of N. marilynae

do reach 32.2 mm in aquaria (Weitzman and Cobb,

1975). Nannostomus limatus appears to have a proportionately larger eye than N. marilynae, 90% to

100% of the interorbital width versus 70% to 90%.

Thus AT. limatus differs from N. marilynae in adult

size, lack of an adipose fin, more gill rakers, a

larger eye, no second row dentary teeth, and in certain details of the preserved color pattern such as

a black distal edge on the ventral portions of the

anal fin and a distinct dark, narrow edge on the

dorsal and ventral borders of the caudal fin.

Sometimes preserved specimens of N. limatus

show weakly developed horizontal tertiary stripes

(Figure 7). In some cases it might be possible to

confuse preserved specimens of N. limatus with

those of N. beckfordi, especially since both species

lack an adipose fin and have similar scale counts

and are found in the same region of the Amazon

basin. Specimens of N. limatus are here compared

with specimens of N. beckfordi from the Rio Inhangpi at Inhangapi, Para, Brazil, USNM 216547.

Males of N. beckfordi (Figure 9) have a very poorly

developed secondary horizontal stripe, whereas in

N. limatus (Figure 6) this stripe is well pigmented

and formed as a series of about 15 dark spots, one

at each of the anterior bases of the 15 anterior

scales of the second longitudinal scale row (Figure

6). Also males of N. beckfordi have the primary

horizontal stripe broad compared to that of males

of N. limatus, while the pale stripe above the primary stripe in N. beckfordi is nearly nonexistent,

but is well developed in N. limatus (compare Figures 6 and 9).

Females of N. limatus and N. beckfordi are more

difficult to distinguish (compare Figures 7 and 10)

since both have relatively narrow primary horizontal stripes and well-developed pale stripes dorsal to

the primary stripe. As in the male, however, the

secondary horizontal stripe of the female AT. limatus consists of a spot on the anterior field of each

of the anterior 15 scales of the second horizontal

scale row, and this is nearly absent in N. beckfordi.

Nannostomus limatus further differs from N.

beckfordi in having no posterior row teeth in the

dentary (there are 2 to 6 conical posterior row teeth

in N. beckfordi), a different gill-raker count, 11 to

12 + 17 to 18 in N. limatus and 9 to 10 + 17 to 18

in N. beckfordi.

Perhaps N. limatus is related to N. marilynae;

however, in view of the apparent lability of the

characters used here and by Weitzman and Cobb

(1975) and Weitzman (1966) to typologically distinguish the species of Nannostomus, I can form

no testable hypothesis about the phylogenetic relaFI'CI.RE 9.—Nannostomus beckfordi Giinther, USNM 216553, male, 24.9 mm SL,. Rio

Inhangapi at Inhangapi, Para, Brazil.
NUMBER 263 13

FIGURE 10.—Nannostomus beckfordi Giinther, USNM 216553, female, 20.1 mm SL,

Rio Inhangapi at Inhangapi, Para, Brazil.

tionships of N. limatus. Perhaps live colors would

be useful, but so far as I am aware, the fish has

not appeared in the aquarium trade and its live

colors remain unknown.

Discussion

Weitzman and Cobb (1975) were unable to propose what they believed might be reasonable (testable) hypotheses about the relationships among the

species of Nannostomus except that on the basis of

morphology including color pattern, N. marginatus

and N. trifasciatus may be most closely related to

each other and N. eques and Ar. unifasciatus may

be more closely related to each other than to any

other species. The discovery and description of

three new species has shed little new light on the

relationships of the previously known species of

Nannostomus. In fact the discovery of one of them,

N. britskii, has allowed the proposal of a hypothesis

of relationship in conflict with a suggested sister

group relationship between N. eques and N. unifasciatus. On the basis of a complicated synapomorphic color pattern, N. britskii might be considered the closest relative of N. eques. There is no

evidence, however, to suggest that N. britskii is

closely related to N. unifasciatus or that together

N. britskii and N. eques could form a sister group

with N. unifasciatus. It would seem that the liability of character evolution, that is the convergence

and divergence under environmental selection of,

for the most part, relatively simple observed morphological characters discussed above has obscured

many of the phylogenetic relationships of the

species of Nannostomus.
Literature Cited

Weitzman, S. H. Weitzman, S. H., and J. S. Cobb

1966. Review of South American Characid Fishes of Sub- 1975. A Revision of the South American Fishes of the

tribe Nannostomina. Proceedings of the United Genus Nannostornus Giinther (Family LebiasiniStates National Museum, 119(3538): 56 pages, 12 dae). Smithsonian Contributions to Zoology, 186:

figures. 36 pages, 34 figures.

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