A Revision of the

South American Fishes of the

Genus Nannostomus Giinther

(Family Lebiasinidae)

STANLEY H. WEITZMAN

and

J. STANLEY COBB

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 186
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 186

A Revision of the

South American Fishes of the

Genus Mannostomus Gtinther

(Family Lebiasinidae)

Stanley H. Weitzman

and J. Stanley Cobb

SMITHSONIAN INSTITUTION PRESS

City of Washington

1975
ABSTRACT

Weitzman, Stanley H., and J. Stanley Cobb. A Revision of the South American

Fishes of the Genus Nannostomus Giinther (Family Lebiasinidae). Smithsonian

Contributions to Zoology, number 186, 36 pages, 34 figures, 1975.—Based on

newly collected material, this study is a supplemental treatment to an earlier

review of the South American genus Nannostomus by the senior author. Eleven

species are here recognized, N. espei, N. digrammus, N. harrisoni, N. beckfordi,

N. bifasciatus, N. minimus, N. marginatus, N. trifasciatus, N. marilynae (new),

N. unifasciatus, and N. eques. An artificial key to all these species is included.

Additional collections of two species, N. bifasciatus and N. minimus, have allowed

a more complete description of the former and recognition of the latter, which

had been listed as a synonym of N. beckfordi by Weitzman (1966). One new

species, N. marilynae, is described from the Amazon basin. The interrelationships

of all the species are reconsidered and because the new material has revealed

intermediate conditions in the characters formerly used to separate the nominal

genera Nannostomus and Poecilobrycon, the latter is here regarded as a synonym

of the former. Relationships among the species are not clear from the population

samples currently available for study, but it seems probable that N. espei, in the

aggregate of its characters, is the most primitive species in Nannostomtis and not

closely related to any other species. N. marginatus and N. trifasciatus, undoubtedly closely related to each other, are not particularly close to any other species

or species group. The same may be said for the related N. eques and N. unifasciatus in their relation to other species. The remaining species all seem distinct

from one another and their possible relationships to other species in the genus

remain obscure.

Evolutionary trends toward specialization within the genus appear to be development of longitudinal stripes, development of oblique bands, loss of ossification

of the sensory canal in the second infraorbital bone, an elongation and thickening of the anal-fin rays of males as an aid to fertilizing eggs, possibly the development of an ocellus in the dorsal lobe of the caudal fin, and development of

nuptial tubercles on the ventral surface of the head. Thickening of anal-fin rays

occurs to a certain extent in other lebiasinids, but it is carried to an extreme

in some species of Nannostomus. Nuptial tubercles occur in another lebiasinid

genus, Pyrrhuh'na, but their high concentration on the ventral surface of the

head in Ar. unifasciatus is a specialization not found in Pyrrhulina.

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

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The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Weitzman, Stanley H.

A revision of the South American fishes of the genus Nannostomus Giinther (family Lebiasinidae)

(Smithsonian contributions to zoology, no. 186)

Supt. of Docs, no.: SI 27:186

1. Nannostomus. 2. Fishes—South America. I. Cobb, J. Stanley, joint author. II. Title. III.

Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 186.

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Contents

Page

Introduction . 1

Classification of the Family Lebiasinidae . 2

Genus Nannostomus Giinther 3

Artificial Key to the Species of Nannostomus . 6

Nannostomus espei (Meinken) 7

Nannostomus digrammus Fowler 9

Nannostomus harrisoni (Eigenmann) 11

Nannostomus beckfordi Giinther 14

Nannostomus bifasciatus Hoedeman 16

Nannostomus minimus Eigenmann 21

Nannostomus marginatus Eigenmann . 25

Nannostomus trifasciatus Steindachner 27

Nannostomus marilynae, new species 29

Nannostomus unifasciatus Steindachner 33

Nannostomus eques Steindachner 33

Sumario . 35

Literature Cited 36

A Revision of the

South American Fishes of the

Genus Nannostomus Giinther

(Family Lebiasinidae)

Stanley H. Weitzman

and J. Stanley Cobb

Introduction

Since publication of a review (Weitzman, 1966)

of the nominal lebiasinid genera Nannostomus and

Poecilobrycon, the senior author has accumulated

specimens of several of the known species from new

localities, obtained specimens that show unrecorded variation, and received additional specimens of two poorly known species, N. bifasciatus

Hoedeman and N. minimus Eigenmann. Examples

of a new species from Brazil, N. marilynae, have

been in the senior author's care for nearly 20 years

but were not described as new previously because

all specimens at hand were juveniles. In 1969 one

of us (Weitzman) found live specimens of this new

species in an aquarium shop in the Washington,

D. C, metropolitan area. At the same time two

males of N. digrammus Fowler were obtained, and

life colors of this species are recorded here. All this

new material has allowed a reevaluation of the

generic problem within this group of lebiasinids

as well as a new consideration of species relationships.

Stanley H. Weitzman, Division of Fishes, Department of

Vertebrate Zoology, National Museum of Natural History,

Smithsonian Institution, Washington, D.C. 20560. J. Stanley

Cobb, Department of Zoology, University of Rhode Island,

Kingston, Rhode Island 02881.

To help the aquarist identify these popular

aquarium fishes, as well as to illustrate the differences between the color patterns of live and preserved specimens, we have included drawings as

well as photographs of preserved specimens where

appropriate. In addition we have included available photographs of live specimens even though

most of these are without locality information.

The key of Weitzman (1966) has been revised

to accommodate a new generic concept, to admit

the new species, to include N. minimus, which

was formerly considered a synonym of N. beckfordi, and to correct an error. The species names

N. beckfordi and N. digrammus had been inadvertently exchanged in the original key.

The methods for taking counts and measurements and the terminology used here for the color

pattern of Nannostomus species are the same as

those outlined by Weitzman (1966). In brief the

term stripe is restricted to elongate pigmented

areas that extend horizontally along the sides of

the fish. The primary stripe is the main dark midside stripe usually extending from the snout, across

the eye, opercle, and body to the caudal-fin base,

and sometimes onto the caudal fin. The dark

secondary stripe is dorsal to the primary stripe and

lies between the dark pigment of the back and the

pale stripe just dorsal to the primary stripe. The

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

dark tertiary stripe is the ventralmost and may

extend from the lower jaw, across the ventral part

of the operculum, ventral to the base of the pectoral fin, across the ventrolateral part of the belly

just dorsal to the base of the pelvic fins, and to

the base of the anal fin. The nocturnal oblique

bands are areas of pigment, usually two on each

side, that may be present in the pale stripe above

the primary stripe, and in the primary stripe itself. The oblique bands are usually present in

preserved specimens. In life they are weak or absent

in daylight but very dark at night when the horizontal stripes are often very pale. Females sometimes show nocturnal color while breeding. Permanent blotches occur only in one species, espei,

as large pigment areas (see Figures 1 and 2) that

do not fade during the day; the borders of these

blotches remain dark under all circumstances. In

preserved specimens (Figure 1) or in specimens

subordinate to others in agonistic contexts or

specimens responding to gross environmental disturbances (Figure 2), the center of the blotch may

almost completely fade away.

Many people have contributed to the completion

of this study. We are most grateful to the following

persons who have graciously loaned specimens in

their care: James E. Bohlke, Academy of Natural

Sciences, Philadelphia (ANSP); Myvanwy Dick,

Museum of Comparative Zoology, Harvard (MCZ);

William Eschmeyer, Warren Freihofer, and Pearl

Sonoda, California Academy of Sciences (CAS);

P. H. Greenwood, British Museum of Natural

History (BMNH); H. Nijssen, Institute of Taxonomic Zoology (Zoological Museum), University

of Amsterdam (ZMA); Donn Rosen, American

Museum of Natural History (AMNH); J. Gery

(private collection); and Loren P. Woods, Field

Museum of Natural History (FMNH). Susan

Karnella, Edgar N. Gramblin, Bruce Hodess, and

Robert Kanazawa have aided in the curation of

specimens in the National Museum of Natural

History (formerly under the United States National Museum: USNM). All photographs are by

S. Weitzman and William L. Fink, with considerable help from Robert Kanazawa. Marilyn Weitzman provided typing and technical assistance. All

drawings are by the senior author except Figure

14, which is by Marion Dalen.

Ross Socolof and, through him, Louis Chung

provided live specimens with locality information

of N. espei and N. minimus.

James E. Bohlke, Robert H. Gibbs, Jr., William

L. Fink, and David W. Dunham read the manuscript and offered valuable suggestions.

The work was partially supported by Smithsonian Research Award 235340 to the senior

author and a Smithsonian postdoctoral fellowship

to the junior author.

Classification of the Family Lebiasinidae

We wish to recognize in a formal way a change

in characoid classification since the publication of

the senior author's papers on lebiasinid fishes in

1964 and 1966. Greenwood, Rosen, Weitzman, and

Myers (1966) recognized several families of characoids, one of these being the Lebiasinidae. Weitzman (1964 and 1966) considered the Lebiasinidae

as a subfamily of the inclusive family Characidae,

which equals the Characoidei of Greenwood, et

al. (1966). We believe that acceptance of several

families of characoids is a realistic expression of

the amount of divergent evolution that has occurred in the group and therefore adopt the following arrangement for the lebiasinid fishes based

on the osteological work of Weitzman (1964). Definitions of the groups below, except for the genera,

can be found in the works of Weitzman (1964 and

1966). In the earlier papers the subfamily Lebiasininae is equivalent of the Lebiasinidae in the

present paper, while the former tribes Lebiasinini

and Pyrrhulinini now become the Lebiasininae

and Pyrrhulininae, respectively. The former subtribes Pyrrhulinina and Nannostomina now become the tribes Pyrrhulinini and Nannostomini.

Family Lebiasinidae Eigenmann (1910)

Subfamily Lebiasininae Eigenmann (1910)

Genus Lebiasina Valenciennes (1846), in Cuvier and

Valenciennes (1846)

" Piabucina Valenciennes (1849), in Cuvier and

Valenciennes (1849)

Subfamily Pyrrhulininae Eigenmann (1910)

Tribe Pyrrhulinini Eigenmann (1910)

Genus Pyrrhulina Valenciennes (1846), in Cuvier and

Valenciennes (1846)

" Copeina Fowler (1906)

" Copella Myers (1956)

Tribe Nannostomini Eigenmann (1909)

Genus Nannostomus Giinther (1872)
NUMBER 186

Genus Nannostotnus Giinther

There are four nominal genera in the tribe Nannostomini: Nannostomus Giinther (1872), Poecilobrycon Eigenmann (1909), Archicheir Eigenmann (1909), and Nannobrycon Hoedeman (1950).

Only Nannostomus is recognized here, and an

explanation of this involves a discussion of the

possible relationships among the following species,

all those known in the genus. The order of the

species in this list does not constitute an interpretation of phylogenetic relationships but follows

the order given in the artificial key below and the

order of treatment here.

Nannostomus espei (Meinken), 1956

Nannostomus digrammus Fowler, 1913

Nannostomus harrisoni (Eigenmann), 1909

Nannostomus beckfordi Giinther, 1872

Nannostomus bifasciatus Hoedeman, 1953

Nannostomus minimus Eigenmann, 1909

Nannostomus marginatus Eigenmann, 1909

Nannostomus trifasciatus Steindachner, 1876

Nannostomus marilynae, new species

Nannostomus unifasciatus Steindachner, 1876

Nannostomus eques Steindachner, 1876

Weitzman (1966) accepted two generic level

taxa in the Nannostomini, Nannostomus and Poecilobrycon. Nannobrycon was included as a subgenus of Poecilobrycon, and Archicheir, based on

juveniles of the type-species of the genus Poecilobrycon (P. harrisoni), was listed as a synonym of

Poecilobrycon (compare Figure 8 with fig. 5 in

pi. 37 of Eigenmann, 1912). Poecilobrycon was recognized by Weitzman (1966) because of its longer

snout and the presence of a bony canal in the

second infraorbital bone for the infraorbital

branch of .the laterosensory canal. Presumably the

presence of a canal in the second infraorbital bone

is primitive for the tribe Nonnostomini since the

canal is found in most characoids as well as in all

other members of the subfamily Lebiasininae. It

is also found in several but not all of the species

in the tribe Pyrrhulinini. There appears to be a

trend within the family Lebiasinidae for a reduction of the sensory canal system in the more specialized genera and species. The species with a

reduced canal are frequently small in adult size.

Thus retention of the canal in the second infraorbital bone as found in N. harrisoni, N. unifasciatus, and N. eques should be primitive for the

genus, and its loss in the remaining species an

advanced character that would unite them in a

specialized genus, Nannostomus, according to the

principles of phylogenetic systematics. However,

one very simple character, and a reductive one at

that, usually cannot be given much importance

in relating fishes.

There is some evidence that the absence of the

canal in the second infraorbital bone may simply

be a function of the size of individual specimens

in a given species. Both N. harrisoni and N. bifasciatus are large species, reaching at least 41.5 and

43.2 mm in standard length, respectively. A canal

is always present in N. harrisoni, but examination

of many collections of N. bifasciatus shows that not

all specimens have a well-developed canal; the

canal is absent in many large specimens as well as

in small ones. Nannostomus unifasciatus is also

fairly large, reaching at least 38.3 mm in standard

length, while N. eques, although usually large at

30 to 33 mm in standard length, has been known

to reach 35.4 mm in standard length (see Weitzman, 1966, and the data recorded in the present

text). Both species have a canal in the second infraorbital bone. Of the remaining species to reach an

intermediate size, old aquarium specimens of N.

beckfordi examined here reach at least 35.0 mm

in standard length, and old aquarium specimens

of N. trifasciatus attain at least 35.3 mm in standard length. These last two species always lack a

canal in the second infraorbital bone as do all the

remaining, smaller species. Thus the smaller

species and some of those of intermediate size, N.

beckfordi and N. trifasciatus, lack the canal in the

second infraorbital bone, whereas the larger species

have it or tend to have it.

Small size is a specialization in this group, and

infraorbital canal loss and small size seem correlated. Although these two specializations appear

to constitute a trend for the genus, the size at

which the canal is lost is not sharply defined, and

at least one species, N. bifasciatus, may or may not

have the canal. Therefore there seems little reason to recognize two genera based on these two

characters.

Of other characters that might bear on the problem of interspecific relationships, color pattern is

at least interesting and sometimes helpful. So far,

N. unifasciatus is the only species reported to have

an ocellus in the upper caudal-fin lobe. Even this

species does not always have the ocellus, it being
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

best developed in specimens from Guyana (see

Weitzman, 1966, and Figures 31 and 32 of this

paper), where adults of both sexes have the dark

spot surrounded by orange and white pigment.

The spot has not been recorded for N. eques and

cannot be seen in many specimens; however,

orange pigment, similar to that of N. unifasciatus,

is often present in the caudal fin of both sexes,

and some specimens (see female in Figure 34) have

a moderately developed ocellus. Young specimens

of N. bifasciatus (Figures 15 and 16) have a

caudal-fin spot similar to that of N. unifasciatus,

but this seems to disappear completely in adults

(Figure 17). Young of N. harrisoni also have a

spot (Figure 8, specimen 18.0 mm in standard

length), but this also disappears or mostly disappears in adults (Figure 7). Most, if not all,

young specimens in all species of Nannostomus

have much dark pigment on the caudal fin (see

Figure 8 of N. harrisoni), and the dark pigment

on the caudal fins of adult N. eques and N. unifasciatus is probably a retention in adults of this

juvenile dark color. The wedge-shaped black pigment on the caudal fins of adult Nannostomus of

most species represents what is left of this dark

juvenile pigment. No ocellus seems to occur in

any of the remaining species of Nannostomus except in one population sample of N. trifasciatus

(Figure 26) and a sample of juvenile N. beckfordi

(see below). The dorsal wedge (Figures 23 and

24) in N. marginatus may possibly be derived from

the same pigment area as the ocellus.

It would seem that N. harrisoni, N. bifasciatus,

N. unifasciatus, and N. eques may be related by

the following tendencies: to have a sensory canal

in the bony substance of the second infraorbital,

to attain a comparatively large size, and to have

an ocellus or at least a dark pigment spot in the

caudal fin of adults. The first two characters are,

in all probability, primitive for the genus and

therefore not useful in relating these species. The

third character, an ocellus or ocellus-like spot, perhaps a specialization for the genus, is also found

in the young of a least one population of N. trifasciatus (Figure 26). Also much black pigment

is present in the caudal fin of juveniles of other

species of Nannostomus. Dr. David Dunham has

shown us color slides of live juveniles of a population sample of N. beckfordi that display an ocelluslike spot. Without developmental studies of caudalfin pigment, probably correlated with behavioral

observations, it would be difficult to say with assurance whether the spot is primitive for the genus

and is being lost in some species or that it is

specialized and indicates a common ancestor for

the species which have a tendency to possess it.

We believe the appearance of this ocellus in N.

trifasciatus and the widespread occurrence of

black pigment in juveniles of most (and perhaps

all) species of Nannostomus indicates a genetic

background within the genus and that therefore

the possession of an ocellus by a few species is at

best a poor indicator of relationships. Its appearance could be convergent and correlated with a

recognition behavior pattern common to many

members in the genus. For example, Weitzman

(1966) reported two color forms of N. marginatus

with somewhat different but strongly patterned

caudal fins whose sharp color contrast might be

recognition marks of some sort. In one form, probably from the lower Amazon basin, the caudal fin

bears a bright silvery area between two black

wedges, one dorsal and one ventral to the silvery

area. The black wedges are extensions of the primary and secondary stripes onto the fin. In a color

form from the Colombian Amazon, the silvery

area is replaced by brilliant cardinal red. These

areas may serve the same behavioral purpose as

the ocellus in some other species of Nannostomus.

It would seem safe to say that N. eques and N.

unifasciatus are related because of their shared

specialized caudal fins and correlated specialized

swimming position (Weitzman, 1966). Of these

two species N. unifasciatus may be the most primitive in overall color pattern, and it certainly has

a color pattern most similar to that of the other

large species, N. harrisoni and N. bifasciatus (see

discussion below). Nannostomus eques is unique

because it has more horizontal stripes, as many

as five, than the other species, which have one,

two, or three.

It is difficult to relate N. harrisoni and N. bifasciatus to the other large species on the basis of

color pattern. Although the color patterns of N.

unifasciatus, N. harrisoni, and N. bifasciatus are

at first glance similar, N. bifasciatus has a much

better developed secondary horizontal stripe than

the other two, which have the stripe nearly absent

or very pale on both preserved and live specimens.

In live specimens of N. harrisoni the posterior half
NUMBER 186

o£ the primary horizontal stripe becomes very dark

in courting males, and the anterior half fades to

the point of disappearing. This may be unique for

the species, as we have not observed this change in

any other species of Nannostomus (but we have

not seen live N. bifasciatus). Preserved specimens

of N. bifasciatus further differ from those of N.

harrisoni in having the posterior oblique band

anterior and dorsal to the anal fin, whereas those

of N. harrisoni have the band extending anterior,

dorsal, and posterior to the anal fin. The posterior

oblique band is not evident in preserved or live

specimens of N. unifasciatus. Thus of the species

N. unifasciatus, N. bifasciatus, and N. harrisoni,

there seems little to relate them on the basis of

stripes and bands. All have these typical components found in the Nannostomus color pattern but

each is specialized in its own way in regard to

details.

All the large species, N. unifasciatus, N. eques,

N. bifasciatus, and N. harrisoni, have orange to

reddish pigment associated with the nares and

caudal fin in life, but N. harrisoni males also have

this type of pigment on the anal and pelvic fins,

and males and females of N. eques have orange

pigment on the anal fin. In N. eques the narial

pigment is usually gold to orange (rather than the

red of the other species). Again all these species

have xanthic pigments in more or less similar

areas but each is somewhat different from the

others.

Red, orange, or gold pigments are not limited

to the four species described above. They are also

found in at least N. beckfordi, N. marginatus, N.

trifasciatus, N. digrammus, N. marilynae, N. minimus, and N. espei. These have some red, orange,

gold, or yellow pigment in the area of the caudal

fin, anal fin, and/or the nostrils. In N. digrammus,

N. marilymae, N. minimus, and N. espei this pigment is gold or pale orange and sometimes absent.

It is red or orange in the other species. Welldeveloped secondary horizontal stripes occur in

N. marginatus, N. trifasciatus, N. bifasciatus, and

N. eques but nowhere else. Nannostomus marilynae may at times have a moderately developed

secondary stripe, and weak secondary stripes may

occur in N. beckfordi and especially in N. digrammus. The secondary stripe of N. minimus is very

poorly developed. The various pigment patterns

in the species of Nannostomus are difficult to

interpret in terms of relationships and, like morphometric and meristic characters in this group,

may be evolutionarily so labile that no interpretative confidence can be placed in them.

In view of the interpretative difficulties it seems

advisable to recognize only one genus, Nannostomus Gunther (1872), of which Poecilobrycon

Eigenmann (1909) and Archicheir Eigenmann

(1909) are synonyms (see synonymies of Weitzman,

1966). Nannobrycon Hoedeman (1950) could be

recognized for N. eques (the generic type) and N.

unifasciatus because of their unique caudal-fin

structure and "slant" swimming habit. To us, Nannobrycon hardly seems worth recognizing until

studies on the behavior and anatomy associated

with this singular habit are accomplished. In this

connection, although the new species described

here, N. marilynae, swims horizontally, the female

in Figure 29 has the lower lobe of the caudal fin

larger than the upper, similar to N. eques and

N. unifasciatus. The distal end of one of the rays

of the upper lobe has shifted ventrally and lies

between the upper and lower lobes, approaching

the condition found in the homologous rays of N.

eques and N. unifasciatus. The caudal fin of the

male (same figure) appears similar to the majority

of the species of Nannostomus.

Two other characters need some discussion. Both

N. minimus and N. digrammus have greatly modified anal fins in the male and might be considered

related because of this shared specialization

(compare Figures 3 and 18). This could be convergent; these species do not appear closely related

on any other basis and the developement of the

anal fin appears somewhat variable in N. digrammus. Thickened anal-fin rays are found in several

species of Nannostomus (see Weitzman, 1966) and

such rays are also present in the subfamily Lebiasininae as well as in the tribe Pyrrhulinini of the

subfamily Pyrrhulininae. Thus the presence of

thickened anal-fin rays and a trend toward a modification of the anal fin is characteristic of the

family. This greater degree of specialization in

the anal fin of two species of Nannostomus could

be convergent.

A few of the species of Nannostomus are known

to have contact organs, hypertrophic dermal cells

in the form of tubercles. Nannostomus bifasciatus

and Ar. unifasciatus were reported to have such

tubercles by Wiley and Collette (1970). Nannosto-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mus minimus (see below) also has these tubercles.

No tubercles were found in other species but if

they hadn't appeared in the photograph of a live

male, N. minimus (Figure 21), we would have

missed them in our examination of preserved

specimens of this species. In view of these facts we

are not prepared to say that these structures are

absent in other species of Nannostomus, especially

since similar tubercles are found in other lebiasinids, (tribe Pyrrhulinini, see Wiley and Collette,

1970). We can draw no conclusions regarding

relationships of the species based on presence or

absence of tubercles.

Despite the fact that no really useful or phylogenetically significant characters that would warrant

recognition of two genera in the tribe Nannostomini

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 lobe 2

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

lobe 10

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) Nannostomus 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 intensifiied 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 7

4. Adipose fin always absent 6

Adipose fin always present 5

5. Snout in eye 1.1 to 1.3. Total vertebrae 34 or 35. Sensory canal absent in second infraorbital

bone. Secondary horizontal stripe usually well developed Nannostomus digrammus

Snout in eye 0.9 to 1.0. Total vertebrae 38 or 39 in adults. Sensory canal present in second

infraorbital bone. Secondary horizontal stripe usually poorly developed

Nannostomus harrisoni

6. 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

7. Anal fin of males short, not reaching to caudal fin when adpressed to body 8

Anal fin of males long, reaching caudal fin when adpressed to body. Nannostomus minimus

8. Scales in a lateral series 24 to 27. Adipose fin present (sometimes absent in N. trifasciatus) ... 9

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

9. Oblique (vertical) bands broad; anterior band extending posteriorlly 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 Nannostomus trifasciatus

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 Nannostomus marilynae

10. 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 unifasdatus

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
NUMBER 186

have been found in the present study, we do

believe that there are some indications of relationships among the species. Certainly N. eques and

N. unifasciatus seem close because of their swimming behavior and correlated caudal-fin modification, even though N. marilynae may occasionally

show a bit of the same anatomical modification.

Weitzman (1966) noted that N. marginatus and

N. trifasciatus appear close because of very similar

and probably specialized color patterns and we see

no reason to alter this view. The other species, N.

espei, N. beckfordi, N. digrammus, N. harrisoni,

N. bifasciatus, N. marilynae, and N. minimus,

seem to stand alone and we cannot relate them to

other species with confidence.

On the basis of the analysis by Weitzman (1966)

and the discussion above, we are unable to state

with certainty which species or group of species

in the genus Nannostomus may be most primitive.

The presence of a complete, well-formed bony

sensory canal in the second infraorbital bone is

obviously primitive. The subtle body shape in N.

espei (suggesting that of Pyrrhulina, see discussion

of N. espei by Weitzman, 1966) may be primitive,

whereas the more cylindrical shape of some other

Nannostomus species, for example N. harrisoni,

may be more specialized. The profile of the scales

and body shape of N. espei are suggestive of those

in Pyrrhulina, and the black-spot color pattern of

N. espei is similar to that of Pyrrhulina vittata

Regan and Pyrrhulina spilota Weitzman. Nannostomus espei appears to lack oblique bands and

only weakly shows horizontal stripes, the former

being absent in Pyrrhulina, Copeina, and Copella,

while the latter (what may be equivalent to a primary horizontal stripe) is developed to a certain

extent in some species of Pyrrhulina and Copella,

for example Copella nattereri Steindachner and

Copella metae Eigenmann. This suggests that in

the aggregate of its characters N. espei is the most

primitive species, even though it lacks a canal in

the second infraorbital bone. However, both the

body and scale characteristics of N. espei are very

subtle, and the color pattern similarities between

N. espei, C. vittata, and C. spilota, although far

from being subtle, could be convergent. Possibly

N. espei is on a separate line of evolution from the

other known species of Nannostomus, retaining

several primitive characters and perhaps losing the

second infraorbital bone independently.

We propose that the ancestor of Nannostomus

probably had a body and scale shape somewhat

approaching that of Pyrrhylina, that it had only

the primary longitudinal stripe (probably poorly

developed), lacked the oblique bands, had only

a slightly modified anal fin in the male (somewhat

thickened but not elongate rays), had an unmodified caudal fin, no caudal-fin ocellus, little orange

or red pigment, and probably not much black

caudal-fin pigment.

Nannostomus espei (Meinken)

FIGURES 1, 2

DIAGNOSIS.—Secondary and tertiary horizontal

stripes absent; primary horizontal stripe poorly

developed; permanent blotches on sides of living

specimens (in life blotches do not disappear during

daytime). Nannostomus espei differs from all other

species of Nannostomus except N. marginatus in

having only 22 scales in a lateral series (N. minimus has 23 or 24). Caudal fin with 10 principal

rays terminating in dorsal lobe and 9 in ventral

lobe.

DISCUSSION.—Mr. Louis Chung has very kindly

provided us with information on the discovery and

geographical distribution of N. espei. Mr. Chung

first found the fish in 1953 in the "upper Kurupung River, a tributary of the Mazaruni River."

From five expeditions during 1953 and 1954 he

found N. espei to be distributed in the Mazaruni

River from Peaima Falls to Toboku Falls and in

the Kurupung, Eping, and Tabuba rivers, all tributaries of the Mazaruni River, Guyana. He reports that they live in water with a pH of 5.8.

We have no further comments or information

on this species supplemental to the description by

Weitzman (1966). Recent notes on aquarium care

can be found in Sterba (1970).

SPECIMENS EXAMINED.—2, USNM 210704, SL $

25.0 mm, 9 27.2 mm, specimens received alive from

Mr. Ross Socolof, who received them from Mr.

Louis Chung, an aquarium fish exporter-importer

in Georgetown, Guyana. (Mr. Ghung informs us

that he collected these specimens of N. espei from

the Eping River, tributary to the Mazaruni

River, Guyana, 1973. These specimens appear in

the photograph, Figure 2.)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURK 1.—Xannostomus espei, SU 50214 (now at CAS), male, 27.6 mm SL,

Mazaruni River system, Guyana, 1955.

FIGURE 2.—Live Xannostomus espei, USNM 21074, female above, 272 mm SL, male below, 25.0

mm SL, Eping River, tributary to the Mazaruni River, Guyana, 1973.
NUMBER 186

Nannostomus digrammus Fowler

FIGURES 3, 4, 5

DIAGNOSIS.—Primary horizontal stripe always,

and secondary usually, well developed. Tertiary

horizontal stripe absent. Nocturnal oblique bands

present, permanent blotches absent. Adipose fin

always present. Snout in eye 1.1 to 1.3. Total number of vertebrae 34 or 35. Anal fin of male elongate, reaching caudal fin when adpressed to body.

Caudal fin with 10 rays terminating in dorsal lobe

and 9 rays in ventral lobe.

DISCUSSION.—The synonymy remains the same

as recorded by Weitzman (1966). Sterba (1970)

appears to have correctly identified this fish.

The list of specimens examined consists of new

geographical localities recorded for this species.

Most of these localities are adjacent to those recorded by Weitzman (1966), but one extends the

known range in the Rio Madeira up into the Rio

Guapore between Brazil and Bolivia.

The Rio Madeira-Rio Guapore collection consists of 31 individuals of both sexes; all are relatively small, not exceeding 22.9 mm in standard

length. The species attains a large size, for old

aquarium males from unknown localities reach a

length of 29.0 mm in standard length. The mature

adult males of the Guapor£ population sample

have smaller anal fins (the rays are shorter in

length) than specimens of similar size from the Rio

Negro around Manaos, from the Rio Branco system in Brazil, or from the Rupununi system in

Guyana. Meristic and morphometric data taken

on the specimens examined here mostly fall within

the ranges of the specimens previously recorded

by Weitzman (1966). A few specimens from the

Rio Purus were found with one less scale in a

lateral series (range 24 to 26 instead of 25 or 26).

Fin-ray counts and the basic tooth pattern remain

as given by Weitzman (1966). Morphometric data

show some degree of geographic variation, but

the relatively few specimens from most localities

and the irregular distribution of these localities

defeat any attempt at meaningful analysis.

There seems to be some confusion in the aquarium literature regarding the identification of N.

digrammus and N. beckfordi. Under the name

Nannostomus beckfordi beckfordi Giinther, Sterba

(1962:209) described what he called a "Golden

pencilfish." As a reference for this account he cited

Meinken (1951:147), who referred to a pencilfish

that he called the "gold-anomalus." Meinken

identified his "gold-anomalus" as Nannostomus

anomalus Steindachner (here considered a synonym of N. beckfordi). Sterba (1970:159) gives

(freely translated) the following description and

life colors for this fish: Back red brown; sides

yellowish to yellow silver; belly white. Blackish to

FK;IRF. 3.—Xntinoslomus digrammus, SV 50248 (now at CAS), male, 24.5 mm SL, Igarape do

Sapucua into Lagoa Sapucua (this lake drains into the Rio Trombetas). Para, Brazil. 7 June 1924.
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 4.—S'annostomus digrammus, SU50250 (now at CAS), male, 20.8 mm SL, Igarapc do Mai

Joana, a tributary of the Rio Xegro near Manaos, Amazonas, Brazil, 25 December 1924.

deep black stripe from tip of snout across eye to

caudal fin. Above this occurs a gold stripe and

above this an iridescent green zone. Scales above

middle of sides dark edged. Fins yellowish to reddish, anal fin and lower lobe of caudal fin beautiful brick red. Female pale compared to male, anal

fin "angular" (rounded in male). In male, posterior half of body reddish at spawning time. "Goldanomalus" differentiated from N. beckfordi by

strong gold stripe and characteristic, principally

downward directed, broadening of dark stripe in

vicinity of operculum.

The senior author has kept several geographical forms of N. beckfordi alive, and some (unfortunately without known locality information) have

had a color pattern similar to this description, with

the males becoming very red on the posterior half

of their body during spawning and courtship

behavior (see male in Figure 13). Presumably the

form described by Sterba (1970) is a geographical

form of Ar. beckfordi. We have never seen a fish

we could identify as N. digrammus with life colors

as described above for the "gold-anomalus."

Previous to 1940 one or possibly more species of

Nannostomus appeared both in Germany and the

United States under the name "gold-anomalus."

Weitzman (1966:26) reported that a specimen was

sent before 1940 by Mr. Frederick Stoye to Dr.

George S. Myers as a "gold-anomalus." At that

time Mr. Stoye was in close contact with German

Fir.iRF. 5.—Live Xannostomus digrammus, I'SNM 208423. male, 28.1 mm SL, locality unknown.
NUMBER 186 11

aquarists, and presumably he had specimens of at

least one of the species (if there was more than

one at that time) being distributed in the aquarium trade as the "gold-anomalus." Mr. Stoye's

specimen is a large male N. digrammns.

It thus appears that at one time or another at

least two species of Nannostomus have been called

"gold-anomalus." We tentatively identify the

species described by Sterba (1970) as one of the

several geographical forms of N. beckfordi (see

Weitzman, 1966, for comments on geographical

polychromatism in N. beckfordi), and that represented by Mr. Stoye's specimen as N. digrammus.

The figure (Tafel 58) of the "gold-anomalus" in

Sterba (1970) is similar to some of the forms of

N. beckfordi from the lower Amazon basin, but

further speculative identification of the species in

this photograph, without preserved specimens, is

pointless.

COLOR IN LIFE.—The color recorded here is from

male specimens from an unknown locality. Dorsal

fin, pectoral fins, and pelvic fins entirely hyaline.

Primary horizontal stripe black, especially dorsal

portion of stripe. Ventral portion of primary stripe

in abdominal region rich brown with some orange

color. Secondary horizontal stripe pale black, lacking olive color of back above. Narrow silvery stripe

between primary and secondary horizontal stripes

with pale specular reflections of mixtures of blue,

green, and purple. This silvery stripe without

other colors and in life no evidence or almost no

evidence of two black oblique bands seen in Figure

4 (compare Figures 4 and 5). Primary black horizontal stripe continuous onto ventral lobe of caudal fin, spreading over above half to three-fourths

of extent of about 8 fin rays. Here black becomes

pale and orange-brown pigment prominent. Belly

below primary horizontal stripe silvery white,

except anterior to anal fin and dorsal to vent

where a considerable amount of orange-brown

pigment occurs. Base of anal fin orange brown,

otherwise hyaline. Caudal fin hyaline except as

described above and at base of silvery stripe be -

tween primary and secondary horizontal stripes

where fin is silvery white. Primary horizontal

stripe on head partly black, with considerable

orange-brown pigment. No bright red spot near

narial opening. Silver pigment suffused with pale

or moderately intense orange occurs around nares.

Dorsum of snout olive brown, with a large number

of small black melanopores. Lower jaw silvery

white. Iris white except for black area of primary

horizontal stripe and a small dorsal black area

near interorbital space. Belly and throat regions

white.

SPECIMENS EXAMINED.—31, USNM 208420, 17.8—

22.9 mm, Brazil-Bolivia border region between

Guajara-Marim Mato Grosso from the Guapore

drainage, 1970, Bernard Von Graeve. 1, USNM

208421, SL 18.2 mm, Brazil, Rio Urubu, 25 miles

from Itcoatiara, State of Amazonas, October 1958,

H. Axelrod and H. Schultz. 9, USNM 208418,

SL 15.8-20.4 mm, Brazil, Rio Purus, Boca de

Tapaua, Nov. 1963, H. Schultz, H. Axelrod, and F.

Terofal, (10 specimens from this locality in the

collection of J. Gery.). 3, USNM 208419, SL

17.0-27.0 mm, Brazil, State of Amazonas, Igarape"

into Lago Grande do Manacapuru, Nov. 1963, H.

Axelrod, H. Schultz and F. Terofal. 2, USNM

208423, SL 24.3-28.1 mm, locality unknown, aquarium specimens (photographed alive, see Figure 5

and color description), 1969.

Nannostomus harrisoni (Eigenmann)

FIGURES 6, 7, 8

DIAGNOSIS.—Primary horizontal stripe well developed. Secondary horizontal stripe usually poorly

developed. Tertiary horizontal stripe absent. Nocturnal oblique bands present, permanent blotches

absent. Adipose fin always present. Snout in eye

0.9 to 1.0. Total number of vertebrae 38 or 39.

Anal fin of male not elongate, not reaching base

of caudal fin. Caudal fin with 10 rays terminating

in dorsal lobe, 9 rays in ventral lobe.

DISCUSSION.—Except for the general discussion

above, we have few comments on this species in

addition to those of Weitzman (1966). The synonymy remains the same. Sterba (1970) correctly

cites this fish. Although the photographs of specimens (Figures 7 and 8) presented here were made

primarily to show developmental color pattern

differences in the caudal fin and the tendency of

the young to have a caudal-fin ocellus that is

vaguely similar to the well-developed ocellus in the

Guyana population of N. unijasciatus, these

photos also show other changes in pigment as

the fish progresses from 14.6 mm standard length

to adult size of approximately 35 to 38 mm stand-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ard length. The most obvious change is that of

the primary horizontal stripe, the pale stripe above

it, and the secondary stripe. In a 14.6 mm standard

length specimen the pale stripe dorsal to the primary stripe has a large amount of black pigment

posteriorly (lower fish, Figure 8). By 18 mm standard length this pigment is nearly gone.

Figure 8 shows the late change of the pectoral

FIGURE 6.—Xannostomus harrisoni, SU 50243 (now at CAS), male, 38.5 mm SL,

Georgetown, Guyana, no date.

FK;IRE 7.—Live Xanno.Uomm harrisoni, I'SNM 204075, female above, 38.2 mm SL, male below,

35.1 mm SL, Guvana, 1970.
NUMBER 186 13

FICURF. 8.—Sannoslotnui harrisoni, I'SNM 204075, ><>ung of adults in Figure 7. above lo below,

21.1, 18.0, 16.4, and 14.6 mm SL.
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fin in N. harrisoni from the fin of a postlarva to

that of an adult. This late development led Eigenmann (1909:46) to describe Archicheir minutus

.(from 1 specimen) as being related to Nannostomus, but with peculiar pectoral fins having "broad,

dermal flaps, with hair like fringes." This description fits the normal pectoral fins of all postlarval

Nannostomus. The type of Archicheir was 26 mm

in total length, a large size for a Nannostomus

with postlarval pectoral fins. Eigenmann (1912:

287) notes that Archicheir is probably based on

a specimen that is abnormal and that the genus

might be a synonym of Poecilobrycon. Weitzman

(1966:37, 41), after studying the holotype of the

type-species of Archicheir and comparing it with

the young of N. harrisoni, identified it with the

young of N. harrisoni. Comparison of Eigenmann's specimen (1912, pi. 37: fig. 5) with the 18

mm specimen of N. harrisoni shown here (Figure

8) shows two nearly identical fishes. The specimen

in Figure 8 (that is 21.1 mm in standard length)

is nearly 25 mm in total length, nearly as long as

the type of Archicheir, but has mature pectoral

fins. Apparently this species develops adult

pectoral-fin morphology at about 20 to 22 mm in

standard length. Unfortunately we have almost no

data on the length of specimens at maturity of the

pectoral fins in species other than N. harrisoni,

but specimens less than 20 mm in standard length

of the other species we have examined have adult

pectoral fins.

Nannostomus beckfordi Giinther

FIGURFS 9, 10, 11, 12, 13

DIAGNOSIS.—Primary horizontal stripe well developed, secondary horizontal stripe poorly developed,

and tertiary horizontal stripe absent. Nocturnal

oblique bands present, permanent blotches absent.

Adipose fin always absent. Gill rakers 9 or 10 + 17

or 18. Caudal fin with 10 principal rays terminating

in the dorsal lobe and 9 in the ventral lobe.

DISCUSSION.—Although we list a moderate number of specimens from new localities, these population samples do not extend the known range of

this species to any appreciable extent. Also they

add nothing to the discussion on geographical

variation by Weitzman (1966).

Figure 11 is of the holotype, a male, never before

illustrated. The specimen has at some time been

dried and is now in poor condition. Figure 12 is

of a pair of live specimens that compare favorably

with specimens from Guyana. Figure 12 is included

here in order that a direct comparison can be

made between photographs of live N. beckfordi

Fir.iRF. 9.—Satinostomus beckfordi, SI' ">0261 (now at CAS), male, 30.0 mm SL, I'tinga Forest

Reserve, Bclem, I'ara, Brazil, September 1944.
NUMBER 186 15

FIGURE 10.—Nannostomus beckfordi, SU 50261 (now at CAS), female, 29.5 mm SL, Utinga Forest

Reserve, Belem, Para, Brazil, September 1944.

and the new species N. marilynae. Figure 13 illustrates a male that compares favorably with specimens known from the lower Amazon around

Belem. See Weitzman (1966) for a discussion of

the differences in geographical forms of this

species.

Weitzman (1966) cited N. minimus as a synonym

of N. beckfordi. Nannostomus minimus is here

treated as a valid species (see "Discussion" under

that species).

SPECIMENS EXAMINED.—1, holotype, BMNH

1871-12-28:10, SL 22.0 mm; Guyana, Demerara.

61, USNM 208261, SL 13.1-28.3 mm; Surinam, Para

district, Berlijn, Para Creek, 14 September 1969,

H. P. Pijpers. 12, USNM 208262, SL 20.4-24.0

mm, Surinam, Para district, 43 km south of Paramaribo, a blackwater creek near 'S Lands Basheheer Headquarters at Zanderij. White sand savanna

belt, 9 April 1969, D. W. Dunham. 2, ZMA 104.

206, SL 16.0-26.8 mm; Surinam, Mastenkreek, Berlijn, Para river system, about 60 km from Paramaribo; H. P. Pijpers, 24 March 1962. 3, ZMA

106.179, SL 22.4-22.8 mm; Surinam, Marowijne

district, Ricanau Creek about 5 km from Moengo

via Albina-Moengo road, 4 June 1966, H. Nijssen. 2, ZMA 104.205, SL 14.6-27.8 mm; Surinam,

11.—Xantiottomus beckfordi, BMNH 1871-12-28:10, male, 22.0 mm SL. holotype.

Demerara. Ciuvana, date of collection unknown.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIOURF 12.—Live Xannostomus beckfordi, male above, 26.3 mm SL, female below, 26.9 mm SL,

no locality data, specimens very much like those known to come from Guyana.

Mooi-Wana Creek along Albina-Moengo road,

Cottica River system, 10 March 1962, H. P.

Pijpers. 1, ZMA 104.207, SL 17.0 mm; Surinam,

Matoekasi Creek along Albina-Moengo Road,

Cottica River system, 14 March 1962, H. P. Pijpers. 35, ZMA 102.015, SL 15.5-24.4 mm; Surinam, Coropine Creek near Republick, October

1956, J. van der Kamp. 221, ZMA 100.514, SL

22.9-27.0 mm; Surinam, Suriname River at village

Berg en Dal, 15 March 1952, Mr. Spoelstra. [These

include the holotype and paratypes of Nannostomus beckfordi surinami Hoedeman. Dr. H. Nijssen

states that the holotype apparently never was separated from the paratypes. It is now impossible to

distinguish the holotype from the other specimens

in the jar.] 25, MCZ 16042, SL 16.7-23.7 mm;

Bra/il Lagoa de Piovideiu ia, Municipio of Aninindeua, Belem, Pani, July 1965, X. Menezes. 5,

MCZ 49360, SL 19.7-21.0 mm; Brazil, Rio Apeu,

Boa Vista, Municipio of Castanhal, State of Para,

July 1965, X. Menc/es. 1 ZMA 105. 325, SL 23.7

mm; Surinam, Carolina ( = Malasiel) Creek, about

10 km south of Zanderij, Para district, 14 December

1966, H. Nijssen.

Nannostomus bifasciatus Hoedeman

FIGURES 14, 15, lfi, 17

Previously Ar. bifasciatus was known from four

specimens in poor condition and some photographs and comments in the aquarium literature

(see Weitzman, 1966). Through the field work of

Dr. H. Nijssen, ZMA, the fine collections recorded

below have been made available and form the

basis of the following redescription of this species.

The description is based on 35 specimens, 25.7 to

43.2 mm in standard length, from the Brokopondo

District of Surinam. The largest recorded specimen (an aquarium fish) was 44.1 mm in standard

length.

DIAGNOSIS.—Primary and secondary horizontal
NUMBER 186 17

FIGURE 13.—Live Nannostomus beckfordi, male, 32.4 mm SL, no locality data, specimen very

much like those known to occur from areas around Belcm, lower Amazon River area, Brazil.

Fir.fRF. 14.— S'antwstotnus bifaschrtus, /.MA 106.144, male, 41.2 mm SI., creek, 12 km south of

Brownsueg, Brokopondo District, Surinam, 23 Noveml>er 1966.
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 15.—Nannostomus bifasciatus, ZMA 106.144, young, 17.1 mm SL, same data as specimen

in Figure 14.

stripes well developed, tertiary horizontal stripe

absent. Nocturnal oblique bands present, permanent blotches absent. Adipose fin always absent.

Gill rakers 8+14 . Caudal fin with 10 principal

rays ending in dorsal lobe 9 in ventral lobe.

DESCRIPTION.—Body elongate, sides somewhat

flattened, and body compressed posteriorly. Greatest body depth slightly anterior to dorsal-fin origin

224 (180-300); least depth of caudal peduncle 098

(090-110); length of caudal peduncle 184 (140-

260). Snout tip to origin of dorsal fin 533 (510-

570); snout tip to origin of anal fin 755 (720-810).

Head conic and snout obtuse in both vertical

and horizontal profiles; top of head gently convex

between eyes. Head 259 (220-310); eye 080 (070-

100); snout 083 (070-090); least width of bony

interorbital 078 (070-090).

Premaxillary with 7 or 8 teeth, each bearing 7

cusps in adults; second from lateralmost cusp

largest. Maxillary with 1 tooth bearing 4 or 5 cusps

(usually 5 in adults); second or third cusp from

dorsalmost cusp largest. In adults dentary with

6 to 8 multicuspid teeth in outer row, anterior 7

with 7 cusps, third from median side of tooth

largest, eighth (most posterior) mulitcuspid tooth

notably smaller than others and with 4 or 5 subequal cusps. Sometimes (in large adults) a simple

conic tooth posterior to eighth multicuspid tooth

of dentary. Inner row of dentary teeth simple,

conic, 13 to 15 in number. Replacement teeth of

outer row of "primary" teeth present.

Gill rakers 6 to 8 + 12 to 14.

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

i.ll; pelvic fin ii,7; principal rays of caudal fin

10/9. Adipose fin absent. Anal fin of male somewhat modified. Anterior ray small and nearly or

completely hidden by basal scale. Second ray

enlarged and its anterior edge expanded and carinate. Entire fin larger than that of females and

individual rays thickened laterally and somewhat

enlarged in anterior to posterior plane. Posterior 1

or 2 rays little expanded. In a folded position, rays

FIGURE 16.—Nannostomus bifasciatus, ZMA 106.159, young, 23.2 mm SL, Tapocripa Creek near

Herman's Village, 1 km north of Brokopondo, Surinam, 25 July 1966.
NUMBER 186 19

take an alternate position to each other in regard

to median plane. However, fin not remarkably

modified as in Ar. digrammus and N. minimus.

Scales belonging to that part of third horizontal

scale row below dorsal fin having posterior field

with 1 to 3 or rarely 4 (usually 2) radial grooves;

dorsal and ventral fields with one anterior field

with 3 or usually 4 radial grooves. Scales in lateral

series 23 to 27, most often 25 or 26. Predorsal scales

10 or 11, usually 11.

Total number of vertebrae 35 to 38, usually 36

or 37 (one with 38).

Wiley and Collette (1970) described breeding

tubercles or contact organs in male N. bifascialus

as without a distinct keratinized cap and occurring

on ventral surface of head, lower jaw, and in

several rows posterior to isthmus. We find these

tubercles very small and white in preserved specimens. Adult males with as many as 18 to 25 along

ventral border of subopercular and interopercular

bones. Up to 18 small tubercles scattered over

exposed surface of each dentary, up to 3 on each

maxillary bone, and as many as 20 to 25 on each

third infraorbital bone. At least 5 or 6 occur on

ventral surface of first and second infraorbital

bones. Total number of tubercles on ventral surface of head nearly 150.

COLOR IN ALCOHOL.—Top of head dark brown,

back light brown. Primary horizontal stripe dark

brown, nearly black, extending from snout tip and

lower jaw onto anterior surface of eye, across lower

border of fifth orbital bone and upper border of

fourth orbital bone to over upper limit of preopercle, spreading across opercle to posterior

bony border. On body, primary horizontal stripe

beginning anteriorly underneath fleshy opercular

Hap and extending through lower half of third

scale row (from top), almost completely through

fourth scale row, and through upper third of fifth

scale row. Primary stripe widening posteriorly to

include all of fourth scale row along caudal peduncle and above anal fin. Primary stripe covers

entire ventral half of caudal peduncle and posteriormost 1 or 2 rays of anal fin. Stripe continues

into dorsal portion of ventral lobe of caudal fin,

including at least 2 ventralmost rays of dorsal lobe

of fin. Black of stripe continues nearly to termination of fin rays. Black may form a wedge-shaped

area or simply a rounded "blotch" as in Figure 17.

Ventral to horizontal stripe, sides, and belly white

with a few irregularly scattered melanophores.

Tertiary stripe absent except for a few malanophores just anterior to pelvic-fin base. Silvery or

white stripe dorsal to primary horizontal stripe,

extending from eye to caudal-fin base. Two areas

of large, scattered melanophores occur on stripe,

forming oblique bands. Anterior band covering 3

to 4 scales anterior to a point just ventral to anterior origin of dorsal fin. Posterior band covering

another 3 to 4 scales in area of silvery stripe. Center

of posterior band at a point dorsal to vent. Just

dorsal to pale or silvery stripe, a narrow secondary

stripe arises just dorsal to termination of dorsal

opercular opening and extends posteriorly through

center of second scale row below dorsal fin. Posterior to dorsal fin, stripe may merge with dark color

of back or continue (as in Figure 14) to dorsalmost surface of caudal peduncle to a point posterior to a vertical from the posterior termination

of the anal fin. All median fins, except as described

FIGURE 17.—Xannostomus bifascialus, /.MA 106.15fi. male, 35.7 mm SL, creek tributary to

Surinam River, 500 km south of Botopasi, Surinam, 19 March 1967.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

above for caudal fin and anal fin, colorless except

for a few scattered melanophores. Pelvic and pectoral fins with melanophores scattered along

borders of rays.

COLOR IN LIFE.—Life colors recorded by Weitzman (1966) were based on the description of

Hoedeman (1954). The present description is

based primarily on color photographs taken of

freshly preserved specimens from Marowijne Creek,

Surinam, collected by H. Nijssen, October 1966.

Back dorsal to secondary dark stripe olive brown.

Secondary dark stripe brown to black. Pale stripe

between primary and secondary stripes pale yellow

or pale green to white. Sides and belly ventral to

primary black stripe white. Area on caudal peduncle dorsal and ventral to primary stripe orange

(perhaps more red in life). A similar orange

color at base of anal fin. Hoedeman (1954) recorded a "reddish flush" on the base of the caudal

and anal fins in living specimens. There is an

orange or red spot associated with the nares on

the Marowijne Creek specimens. Hoedeman (1954)

records that the pelvic fins are tipped with "ice

blue." As would be expected, the blue is absent

in the freshly preserved specimens, being quickly

destroyed by formalin.

DISCUSSION.—We have little to add to the discussion under "Remarks" by Weitzman (1966). Hoedeman (1953 and 1954) reported a vestigial

adipose fin in N. bifasciatus. Weitzman (1966)

found no trace of one in Hoedeman's types and

we found none in any specimen examined here.

It has been noted above that a canal in the second

infraorbital bone may be present or absent. Our

present views concerning the relationships of this

species are discussed under the genus.

SPECIMENS EXAMINED.— (Note, all specimens

from Surinam and collected by H. Nijssen unless

otherwise indicated.) 3, paratypes, ZMA 100.513,

SL 27.2-28.3 mm, (only two specimens could be

measured as snout of one damaged), Berg en Dal,

Surinam River, March 1952, Blij-Dorp Expedition

(not collected by Nijssen). 12, ZMA 106.144, SL

28.9-13.1 mm, Witte Creek, 12 km south from

Brownsweg, Brokopondo District, 23 November

1966. 28, ZMA 106.145, SL 34.5-43.2 mm, Marowijne ( = Gran) Creek about 53.5 km south from

dam at Afobaka, Brokopondo District, 25 May

I%6. 12, ZMA 106.146, SL 23.5-36.8 mm, Zij

Creek ( = Gran Man), 1 km northeast of Dombaai

(=Bendi Watra), Gran Rio, 15 km southwest of

Bjoemoe, May 30, 1967. 11, ZMA 106.147, SL

20.4-42.2 mm, Marowijne Creek, Brokopondo

District, 58 km south of dam at Afobaka, 9 June

1966. 22, ZMA 106.148, SL 24.0-35.4 mm, Dabikwen (=Baling) Creek, Brodobaka, 5 km north of

Afobaka, Brokopondo District, 19 September

1966. 14, ZMA 106.153, SL 32.6-43.2 mm, Marowijne ( = Gran) Creek, Brokopondo District, 60 km

south of dam at Afobaka, 8 June 1966. 7, ZMA

106.154, SL 20.5-37.1 mm, Marowijne ( = Gran)

Creek, 51 km south of dam at Afobaka, Brokopondo District, 27 May 1966. 2, ZMA 106.155,

SL 30.2-37.7 mm, Marowijne ( = Gran) Creek, 50.5

km south of dam at Afobaka, Brokopondo District,

6 May 1966. 5, ZMA 106.156, SL 31.4-38.4 mm,

Kwati Watra Creek, Suriname River, 500 km south

of Botopasi, Brokopondo District, 19 March

1967. 10, ZMA 106.158, SL 34.6-37.4 mm, Makami

( = Makambi) Creek, 8 km south of Brownsweg,

Brokopondo District, 13 September 1966. 3, ZMA

106.159, SL 25.8-34.8 mm, Sara Creek, about 31

km south of Borpje Dam, Brokopondo District,

12 October 1966. 2, ZMA 106.160, SL 30.7-33.1

mm, Sara Creek, about 27 km south of Dorpje

Dam, Brokopondo District, 14 October 1966. 2,

ZMA 106.161, SL 24.1-37.1 mm, Marowijne

( = Gran) Creek, 63 km south of dam at Afobaka,

20 October 1966. 7, ZMA 106.163, SL 25.9-38.6

mm, Parwapa ( = Paba) creek, Suriname River,

2.5 km from Botopasi near Voetoenakaba (=Voetoepkaba), 20 March 1967. 4, ZMA 106.164, SL

25.5-39.9 mm, French Guiana, creek emptying into

Marowijne River just west of Mooisantitabbetje,

3 km north of Stoelmanseiland, 19 April 1967. 4,

ZMA 106.165, SL 25.7-35.1 mm, creek emptying

into Marowijne River, Manbari Val, 6 km north

of Stoelmanseiland, 20 April 1967. 6, ZMA

106.166, SL 22.5-32.3 mm, Maka Creek, emptying

into Lawa River, 10 km south of Stoelmanseiland,

Marowijne District, 21 April 1967. 14, ZMA

106.167, SL 27.9-41.8 mm, Soeakisi Creek, emptying into Tapanahony River, 12 km southwest of

Stoelmanseiland, Marowijne District, 22 April

1967. 4, ZMA 106.168, SL 23.3-29.6 mm, French

Guiana, Kamoloea of Saloea Creek, tributary of

Marowijne River, 9 km southeast of mouth of

Gran Creek (in Marowijne District of Surinam),

24 April 1967. 51, ZMA 105.696, SL 16.9-38.2 mm,

Jenjee Creek emptying into Surinam River 7.5 km
NUMBER 186 21

north of Botopasi, 21 March 1967. 2, ANSP

112253, SL 19.3-22.2 mm, French Guiana, Maroni

Creek (Surinam name) into Gaa Kaba, 21 November 1957, J. Gery. 1, USNM 208266, SL 22.5 mm,

French Guiana, Barcarel Tecun, Ouaqui River,

tributary of Maroni River, 9 September 1971, Eric

Remole. 7, USNM 208265, SI 21.6-25.5 mm, small

stream tributary to Ouaqui River, a tributary of

Maroni River, 5 September 1971, Eric Remole.

7, USNM 208264, SL 16.2-25.2 mm, French Guiana,

Ouaqui River, tributary of Marone, 5 September

1971, Eric Remole.

Nannostomus minimus Eigenmann

FIGURES 18, 19, 20, 21

Nannostomus minimus Eigenmann, 1909:42 [original description; type-locality: Erukin, British Guiana; holotype in

Field Museum of Natural History].—1910:427 [listed]; 1912:

282, pi. 36; fig. 5 [description].

Nannostomus beckfordi (in part) Weitzman, 1966:12 [listed

as a synonym under N. beckfordi, holotype not seen].

Weitzman (1966) considered this species a synonym of N. beckfordi. At that time he had based

his opinion on an examination of a "cotype,"

CAS (IUM) 11691, and on the accounts and photographs given by Eigenmann (1909 and 1912). We

have now examined the holotype, FMNH 52771,

and consider it to represent a valid species. The

holotype (Figure 19) and "cotype" are both females, only the holotype being in good condition.

The tertiary line is faded in these specimens but

can be plainly seen in the holotype (Figure 19).

Discovery of adult males and females (Figure 20)

in the collections at the American Museum of

Natural History and the gift of a live pair from

Mr. Ross Socolof now make redescription of the

species possible.

The "cotype," (SL 16.6 mm) CAS (IUM) 11691,

also from Erukin is in poor condition and very

faded. The presence of a tertiary stripe is not

obvious, only a few faded melanophores are present in the region of the base of the anal fin. This

specimen was compared with a 16 mm standard

length specimen of N. beckfordi from Surinam

(USNM 208261). No differences in eye size or

other morphometric characters were found, and

pigment patterns and meristics were the same. It

is our opinion that this Erukin specimen, a paratype of N. minimus, is a specimen of N. beckfordi.

The following description is based on all 9

known specimens, 16.2-22.8 mm in standard length.

DIAGNOSIS.—Primary stripe present, secondary

stripe absent or, if present, extremely weak. Nocturnal oblique bands present, permanent blotches

absent. Adipose fin absent. Males with a greatly

modified anal fin, with rays larger in diameter and

FIGURE 18.—Xannostomus minimus, USNM 210676, male, 20.9 mm SL, Eping River, tributary

of the Mazaruni River, Guyana, 1973.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 19.—Nannostomus minimus, FMNH 52771, female, 16.2 mm SL, holotype,

Erukin, Guyana, October 1908.

considerably longer than those of females (see

Figures 20 and 21). Caudal fin normal, with 10

principal rays terminating in dorsal lobe and 9 in

ventral lobe.

DESCRIPTION.—Body elongate, cylindrical, with a

somewhat compressed caudal peduncle. Greatest

body depth just anterior to dorsal fin 199 (191-

241). Least depth of caudal peduncle 079 (074-

083); length of caudal peduncle 166 (173-235);

snout tip to anterior origin of anal fin 682 (666-

702).

Head conic, slightly flattened dorsally. Head 273

(251-281); eye diameter 101 (094-107); least width

of bony interorbital 084 (078-089).

Premaxillary with 5 teeth, all 4 cuspid; 2 central

cusps largest and equally large except median

tooth, witli second from lateralmost cusp largest.

Maxillary with 1 tooth bearing 3 cusps, central

Fir.i RF 20.—Xannostomus minimus, I'SNM 209207, male above. 16.8 mm SL, female below, 17.8

mm SI.. "KurapiiMg" (probably equals Rumpling River, a tributary of the Ma/aruni River),

Guvana, 1938.
NUMBER 186 23

cusp largest. Dentary with 5 multicuspid teeth in

outer row, anterior 3 teeth with 5 cusps (sometimes

medialmost cusp very small; second from medial

cusp largest). Fourth tooth with 4 cusps, medial

cusp largest, and fifth with 2 or 3 subequal cusps.

Inner row of dentary teeth simple, conic with 4 or

5 laterally placed teeth. Replacement teeth of

outer row inside row, ventral to both inner and

outer tooth rows, all replacement teeth with fully

developed cusps.

Gill rakers 10 + 13.

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

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

Adipose fin absent. Male anal fin modified in a

manner similar to that of N. digrammus, with rays

somewhat thicker and longer than those of female.

Anterior 3 unbranched rays with about same modifications as similar rays in N. digrammus. First 4

branched rays of male thickened as compared to

those of female (compare Figures 18, 19, 20, and

21), but not nearly as thickened in median plane as

those of N. digrammus (see Weitzman, 1966, fig.

6). Anterior 4 branched rays alternately overlapping

one another in collapsed fin, similar to those of

N. digrammus as described by Weitzman (1966:24).

Inclinator muscles extending bilaterally and distally

onto posterior fin ray far beyond base of anal-fin

rays. Anal fin of male when expanded forms a cup

F. 21.—Live Nanno.stomus minimus, I'SNM 210676, female above, 22.8 mm SI,, male below,

20.9 mm SL, same locality data as in Figure 18. Nfales in Figures 18 and 21 are the same specimen. Note contact organ on ventral surface of male's head (little white spots). In the preserved

specimen these are extremely difficult to locate.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

or becomes spoon shaped, as in N. digrammus.

Distal tips of rays confined to a flat mass of tough

nonexpandable fin tissue. Membranes between fin

rays in central area of fin loose and allow considerable expansion of fin in this area even in preserved

specimens.

Scales belonging to third horizontal scale series

and lying below dorsal fin having anterior field

with 3, sometimes 4, radial grooves. Posterior field

with 1 or 2 radial grooves. Dorsal and ventral

fields each with 1 radial groove. Scales in lateral

series 23 or 24; perforated lateral line scales 1 or

2; scales in median series anterior to dorsal fin 9

or 10.

All specimens with total of 36 vertebrae.

Contact organs present in adult males (see male

in Figure 21) but extremely hard to detect in preserved specimens and no count could be made.

COLOR IN ALCOHOL.—Top of head dark brown.

Primary horizontal stripe dark brown, extending

from tip of snout and lower jaw across eye and

opercle but not across fleshy operclar flap. Primary

stripe continues posteriorly on body from point

internal to opercular flap, across midsides onto

caudal peduncle, but apparently absent on caudal

fin. Primary stripe extends across ventral onefourth of third scale row of body and across dorsal

three-fourths of fourth scale row. Secondary stripe

apparently absent or nearly so. A pale stripe present dorsal to primary horizontal dark stripe; pale

stripe with many scattered melanophores. Body

dorsal to pale stripe light reddish brown. Tertiary

stripe present, consisting of rather loosely scattered

melanophores extending along ventral half of

fifth scale row from pectoral girdle (internal to

opercle) just anterior to base of pectoral fin to

and along base of anal fin. Below primary dark

stripe occurs another white area but without

scattered melanophores. Area of oblique bands

indistinct but may be present in pale stripe covering sixth and seventh scales and in area of pale

stripe dorsal to anal fin. Belly pale, white. Dorsal

fin with scattered melanophores but these more

concentrated anteriorly, anal fin also with scattered

melanophores but these more concentrated in

central area of fin. Pelvic and pectoral fins with

few, scattered melanophores in no particular

pattern.

COLOR IN LIFE.—The color recorded here is from

live specimens of USNM 210676 listed below in

the "Material" section. Back and top of head dark

olive brown. Scale borders of back narrowly outlined with black except for those portions of scale

in area of secondary horizontal stripe. Primary

horizontal stripe black, extending from lower jaw

and snout tip through eye along body sides onto

caudal fin where it ends as a diffuse pale black

blotch, not wedge shaped. Primary horizontal

stripe in both sexes with red chromatophores in

area shared by stripe and anterior oblique bands.

Red color of this area somewhat more intense in

male than female. This color in both sexes very

subdued and not forming a bright red blotch.

Secondary horizontal stripe very weak, really existing only along scale margins that enter area immediately above pale horizontal stripe. Here secondary stripe is represented by an increase in number

of melanophores along anterior and posterior scale

margins. Stripe dorsal to primary longitudinal

stripe pale yellowish green to silvery. Tertiary

horizontal stripe black, dark and intense posteriorly, disappearing anteriorly on belly where dark

melanophores become reduced in number until

finally absent, or nearly absent, at a point below

anterior border of anterior oblique band. Oblique

bands not dark except at night when their melanophores expand. Sides between primary horizontal

stripe and tertiary horizontal stripe silvery white.

Abdomen and throat regions silvery white. Reflective color around nares pale yellow to pale orange.

Dorsal portion of iris of eye above primary horizontal stripe yellow, lower portion below stripe

silvery. A small amount of orange-brown pigment

on caudal-fin base in association with primary

horizontal stripe.

Dorsal, pectoral, pelvic, and caudal fins hyaline

in both sexes except for a few melanophores

scattered along rays. Anal fin of female and especially male with a scattering of melanophores in

central region of fin, forming weakly developed

dark blotch. Caudal-fin base pale yellowish green

to very pale orange brown near termination of

primary and secondary horizontal stripes.

MATERIAL.—1, holotype, FMNH 52771, SL 16.2

mm, Guyana, Erukin, Potaro River, C. H. Eigenmann, October 1908. 1, paratype, FMNH 52772,

SL 18.6 mm, Guyana, Amatuk, Potaro River, C. H.

Eigenmann, October 1908. 3, AMNH 14417, SL

17.4-19.1, Guyana, "Kurapung" (probably equals

Kurupung River, a tributary of Mazaruni River,
NUMBER 186 25

just north of Merume Mts.), collector unknown,

1938. 2, USNM 209207, SL 16.8-17.8 mm, same

data as AMNH 14417. 2, USNM 210676, SL 20.9-

22.8 mm, received alive from Mr. Ross Socolof, who

received them with live specimens of N. espei from

Mr. Louis Chung, an aquarium fish exporterimporter at Georgetown, Guyana. (Mr. Chung

informs us that he collected these specimens from

the Eping River, a tributary of the Mazaruni

River, Guyana, early in 1973.)

Nannostomus marginatus Eigenmann

FIGURES 22, 23, 24

DIAGNOSIS.—Primary, secondary, and tertiary

horizontal stripes all well developed. Nocturnal

oblique bands present, permanent blotches absent. Anal fin of males short, not reaching caudal

fin when adpressed to body. Adipose fin always

absent. Scales in a lateral series 21 to 23. Caudal

fin normal with 10 principal rays terminating in

dorsal lobe and 9 in ventral lobe.

DISCUSSION.—Since the account of this species by

Weitzman (1966), five of the "types" of Cyprinodon amazona Eigenmann have been located in the

Department of Ichthyology, California Academy

of Sciences, by the senior author. These specimens

seem to be all that remain of the 18 specimens used

as "types" in the original description (Eigenmann,

1894). Weitzman (1966:28) noted that the typeseries possibly included both N. marginatus and

N. trifasciatus. The existing five "types" are all

N. marginatus.

As noted by Weitzman (1966), the name N.

trifasciatus Steindachner 1876 predates C. amazona

Eigenmann 1894; however, C. amazona predates

N. marginatus Eigenmann (1909). The name C.

amazona has never before been identified with certainty with N. marginatus, see Weitzman (1966),

Fowler (1954), and Hubbs (1926). Nannostomus

marginatus, on the other hand, is a well-established

name, very common in the aquarium literature and

always used in the ichthyological literature for the

species identified with that name by Weitzman

(1966). Thus the name Nannostomus amazonus

qualifies for suppression by the International Commission of Zoological Nomenclature under the provisions of article 79 in the Code (as revised in the

Bulletin of Zoological Nomenclature, volume 29,

part 4, 29 December 1972). No useful purpose

would be served by selecting a lectotype from the

five existing specimens of Eigenmann's Cyprinodon

amazona and substituting the name Nannostomus

amazonus for Nannostomus marginatus on the basis

FIGURE 22.—Nannostomus marginatus, CAS(IUM) 11700, male, 19.5 mm SL, paratypc, Cluck

Island, Essequibo River, Guyana, 1908.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of priority. The current International Code of

Zoological Nomenclature, article 23 (a,b) (also as

revised in the Bulletin of Zoological Nomenclature,

volume 29, part 4, 29 December 1972) states that

". . . the law of priority is to be used to promote

stability and is not intended to be used to upset a

long established name in its accustomed meaning

through the introduction of an unused name which

is a senior synonym." Thus, Cyprinodon amazona

does not qualify as a senior synonym that must be

used. If any other "types" of C. amazona are ever

found and one or some of them prove to be N.

trifasciatus, one of these should be selected as a

lectotype and Cyprinodon amazona placed in the

synonymy of Nannostomus trifasciatus.

Examination of specimens listed below from

Surinam confirms the belief of Weitzman (1966)

that Hoedeman (1954) was in error in finding

different caudal-fin ray counts between specimens

of N. marginatus from Guyana and Surinam. All

specimens counted here had a principal caudalfin ray count of 10/9. Weitzman (1966) noted

that there are at least three geographically different color forms of this species. We have nothing

else to add to the comments of Weitzman (1966)

concerning this species.

MATERIAL.—5, (types of Cyprinodon amazona),

CAS(IUM) 5083, SL 15.5-17.5 mm, Brazil, "Lower

Amazon", F. C. Hartt. 4, ZMA 105.324, SL 16.2-

20.3 mm, Surinam, Carolina ( = Malasie) Creek,

about 10 km south of Zanderij, 14 December 1966,

H. Nijssen. 43, USNM 208257, SL 14.5-21.6 mm,

Surinam, Para Creek, Berlijn, Para District, 14

September 1969, H. J. Pijpers.

FK.I RK 23.—Xatwostomm marginatus, USNM 208257, male above, 18.") mm SL, juvenile below,

15.0 mm SL, Para Creek, Para District, Surinam, 14 September 1969.
NUMBER 186 27

FK.URF. 24.—Live Nannotfomus marginatus, adult male, 21.6 mm SL, locality unknown.

Nanno8tomu8 trifasciatus Steindachner

FIGURES 25, 26, 27

DIAGNOSIS.—Primary, secondary, and tertiary

horizontal stripes all well developed. Nocturnal

oblique bands present, exceptionally broad,

extending from a vertical line 3 or 4 scale rows

posterior from posterior opercular border to posteriorly well past a vertical line from anterior base

of dorsal fin (see Figure 26), frequently past a

vertical from midbase of dorsal fin. Posterior oblique band extending posteriorly well beyond

posterior end of anal-fin base. Permanent blotches

absent. Eye diameter about 1.0 in interorbital

width. Anal fin of male and female short, not

reaching caudal fin when adpressed to body. Adipose fin present or absent. Scales in a lateral series

24 to 27. Caudal fin with 10 principal rays terminating in dorsal lobe and 9 in ventral lobe.

DISCUSSION.—We have little to add to the account

Fiot'RF. 25.—Xannostomus trifasriatus, SV 36999 (now at CAS), male, 27.") mm SI., Shanslio C'.ano,

Pe\as [Pebas] District, Peru, July or October 1936.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 26.—Xannostomus trifasciatus, MCZ 46040, juvenile, 22.0 mm SL, Rio Apeu, Boa Vista,

Munidpio of Castanhal, Para, Brazil, July 1965.

of this species given by Weitzman (1966). Comments on the nomenclatural problem concerning

Cyprinodon amazona have been given above under

N. marginatus. One interesting population sample

of N. trifasciatus has come to hand since 1966.

This is from the Rio Apeu, Boa Vista, Para,

Brazil. This population, known to us only from

preserved specimens, is like other populations of

N. trifasciatus from Brazil except that all specimens have an ocellus variously developed on the

caudal fin. The specimen in which it is best developed is shown in Figure 26. In the least developed

specimen, a female 26 mm in standard length, the

dark spot is limited to a few rather scattered melaFir.i'Ri. 27.—Live Xantwstonnis trifasciatus, female above, 3">.6 mm SL, male below, 34.8 mm,

locality unknown.
NUMBER 186 29

nophores and could easily be overlooked. Most

specimens show the ocellus reasonably well. Weitzman (1966:30, 31) noted that there are several

distinct populations of N. trifasdatus which can

be distinguished by color at least. The ocellated

form seems to be another.

Figure 26 illustrates another feature of N. trifasciatus that is occasionally confusing to both

aquarists and ichthyologists. When at their darkest

the oblique bands in this species are broad, quite

dark (black in life), and divide the fish into alternate vertical rings of pale and dark color. The

operculum is dark in this species when the two

oblique bands are dark. The preserved specimen

in Figure 26 has the dark bands relatively pale

but well represented, and Figure 27 shows a pair

in life in a phase in which the oblique bands are

almost absent.

MATERIAL.—7, MCZ 46040, SL 20.7-28.1 mm,

Brazil, Rio Apeii, Boa Vista, Municipio of Castanhal, State of Para, July 1965, N. Menezes. 1,

MCZ 46041, SL 28.0 mm, Brazil, Igarap6 at km

29 of the Castanhal-Bel£m road, Municipio of

Santa Isabel, State of Para, July 1965, N.

Menezes. 3, USNM 208260, SL 21.2-26.6 mm,

Brazil, Betem, State of Para, November 1953, H.

Axelrod. 8, USNM 179553, SL 19.1-25.9 mm,

Brazil, Rio Urubu, 25 miles from Itacoatiara,

State of Amazonas, October 1958, H. Schultz and

H. Axelrod. 19, USNM 208263, SL 25.5-31.2 mm,

Brazil-Bolivia border region between GuajaroMirim and Mato Grosso, Guarpor^ drainage, 1970,

B. von Graeve.

Nannostomus marilynae, new species

FIGURES 28, 29, 30

Holotype, CAS (SU) 50238, SL 19.5 mm, Brazil,

Rio Negro, rock pools below rapids, Sao Gabriel,

1 February 1925, Carl Ternetz. 27 (4 alizarin

preparations) paratypes, CAS (SU) 50239, SL 16.1-

21.0 mm, same data as holotype. Specimens not

considered as types: 1, CAS (SU) 50240, SL 18.0

mm, Brazil, Rio Negro, Castanheiro, 15 January

1925, Carl Ternetz. 3, CAS (SU) 50241, SL 17.1-

17.6 mm, Colombian-Brazilian border region,

"sandbank north of Cucuhy," 14 February 1925,

Carl Ternetz. 8 (2 alizarin preparations), USNM

209227, SL 22.4-32.2 mm, aquarium specimens.

The following description (except color and as

otherwise noted in text) is based on the holotype

and paratypes only. [Editor's note: See "Addendum" at end of this section.]

DIAGNOSIS.—Primary, secondary, and tertiary

horizontal stripes present. Nocturnal oblique

FIGURE 28.—Nannostomus marilynae, SU 50238 (now at CAS), 19.5 mm SL, holotype, rock pools

below rapids, Sio Gabriel, Rio Negro, Brazil, 1925.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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. Adipose

fin always present. Scales in lateral series 24 to 27.

Anal fin of males not elongate or greatly modified.

Eye diameter about 0.8 in least bony interorbital

width.

DESCRIPTION.—Body elongate, more or less cylindrical, compressed in region of caudal peduncle.

Greatest body depth at origin of dorsal fin 211 (190-

225); least depth of caudal peduncle 090 (083-097);

length of caudal peduncle 188(167-205); snout tip

to origin of dorsal fin 524(510-560); snout tip to

origin of anal fin 757 (739-784).

Head elongate, conic, snout obtuse, especially

in dorsal profile. Head 266(244-286); eye 101(93-

109); least width of bony interorbital 080 (074-089).

Premaxillary with 5 teeth, all 4 cuspid except

the most posterolateral with 5 cusps; second from

anteromedian cusp largest. Maxillary with 1 tooth

bearing 5 cusps, dorsalmost cusp largest or next

2 ventral cusps largest. Dentary with 5 multicuspid

teeth in outer row, each bearing 5 cusps except

most posterior with 3 cusps; median cusp largest

in all instances. Replacement teeth of outer row

FIGURE 29.—Live Xannostomus marilynae, L'SNM 209227, female above, 32.1 mm SL, male

below, 25.6 mm SL, locality unknown.
NUMBER 186 31

present, ventral to outer and inner rows. Inner

row of dentary teeth simple, conic with 6 to 9

teeth placed along entire inner margin of jaw but

more concentrated laterally where 3 to 4 teeth

occur relatively close together.

Gill rakers 8 or 9 + 13 or 14, usually a total of

21 or 22.

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

i,ll; pelvic fin ii,7; principal caudal-fin rays

10/9. Adipose fin always present. Distal tips of

all fin elements of first 10 principal caudal-fin rays

ending in dorsal lobe of caudal fin in nearly all

specimens. Ventral caudal-fin lobe containing all

distal elements of 9 ventral principal rays. In one

female (upper specimen in Figure 29) tenth ray

down from top (dorsal edge of fin) in exact middle

between two lobes; ventral lobe of caudal fin larger

than dorsal lobe; morphology of caudal fin appearing somewhat intermediate between that of N.

eques and N. unifasciatus on one hand and

remainder of known species of Nannostomus on

other. Anal fin of males somewhat modified, anterior 5 or 6 branched rays longer and more expanded in sagittal plane than remaining posterior

branched rays. Compare shape of fin in male and

female in Figure 29. Anal fin of holotype, Figure

28, typical of that of juvenile.

Scales belonging to third horizontal scale series

and lying ventral to dorsal fin, having anterior

field with usually 3 radial grooves; posterior field

with 1 or sometimes 2 radial grooves; dorsal and

ventral fields each with 1 radial groove. Scales in

lateral series 25 to 26; 2 to 4 perforated lateralline scales.

Total number of vertebrae 34 to 36, usually 34

(18 specimens), occasionally 35 (8 specimens),

and rarely 36 (1 specimen).

COLOR IN ALCOHOL.—Top of head light brown.

Back dorsal to secondary horizontal stripe brown,

somewhat lighter than secondary dark stripe.

Primary horizontal stripe extending from snout

tip posteriorly across lower jaw, over first orbital

bone, through eye and operculum, terminating at

posterior border of bony operculum, but not extending onto fleshy opercular flap. On body this

stripe extends from under fleshy opercular flap

through fourth scale row on body and partially

across third and fifth scale rows. Primary stripe

expands posterior to anal fin to cover half of third

scale row and all scale rows ventral to third scale

row. Primary horizontal stripe extends posteriorly

onto lower lobe of caudal fin. In preserved specimens (see Figure 28) caudal-fin portion appears

as a black triangle. Above primary horizontal

stripe, secondary stripe extends from dorsal border

of operculum posteriorly along second scale series

or row, fading into ground color of back somewhat

anterior to adipose fin. Between primary and

secondary stripes, pale stripe has few scattered

melanophores except in areas of oblique bands,

which occur between seventh and ninth and fourteenth and seventeenth scales in a lateral series of

FIGURE 30.—Live Nannostomus marilynae, LJSNM 209227, female, 24.0 mm SL, locality unknown,

specimen exhibits nocturnal color pattern.
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

third scale row. These bands extend ventrally into

primary horizontal stripe (Figures 28 and 30).

Sides below primary stripe pale yellow (white in

freshly preserved aquarium specimens) with few

scattered melanophores. Oblique bands sometimes

extend below primary stripe into this area (Figures

23 and 30). Tertiary stripe extends along sixth

scale row from posterior termination of operculum

to origin of anal fin, running ventral to base of

pectoral fins and dorsal to base of pelvic fin. Dorsal

fin with scattered melanophores more concentrated

on anterior distal region of fin (sometimes with

almost no melanophores). Pelvic, pectoral, and

anal fins with few melanophores except posterior

ray of anal fin, which may be fairly dark.

COLOR IN LIFE.—The color recorded here is from

male specimens from an unknown locality. Top

of head and back olive brown; scales of back bordered with black. Primary horizontal stripe black,

continuous onto ventral lobe of caudal fin, covering proximal one-third to more than one-half of

5 or 6 dorsalmost rays of that fin lobe. Secondary

stripe black but usually weakly developed. Silvery

stripe between primary and secondary horizontal

stripes with a greenish and pale golden reflectant

color, producing what essentially looks like a

burnished greenish silvery stripe. Oblique bands

not dense black except possibly occasionally at

night (see Figure 30), usually as shown in Figure

29. Sides below primary horizontal stripe on abdomen silvery and belly silvery white. Golden to

orange-brown pigment around nares and on anterior border of eye. No bright red pigment at nares.

Pale orange-brown pigment at caudal-fin base,

ventral to black primary horizontal stripe. Dorsum

of snout and head olive brown, lower jaw silvery

white. Throat region white. Dorsal, anal, pectoral,

and pelvic fins entirely hyaline except for dorsal

fin with black melanophores on basal one-third

of fin rays. Caudal fin hyaline except for area

where black primary horizontal stripe continues

onto fin and an area of white pigment ventral to

that stripe.

ETYMOLOGY.—The name marilynae is derived

from that of the senior author's wife, Marilyn

Sohner Weitzman, who has long shared his appreciation for the delicate beauty of members of the

genus Nannostomxis.

DISCUSSION.—The relationships of this species are

not understood. In the key above, it keys closest

to N. trijasciatus, sharing with that species a short

anal fin; possession of primary, secondary, and

tertiary stripes; an adipose fin; and 25 to 26 scales

in a lateral series. It is very different from N. trijasciatus by its sharply different oblique bands

(compare Figures 26 and 30), life colors (compare

color description in Weitzman, 1966 of N. trifasciatus with that of N. marilynae published here),

and somewhat smaller adult size, not exceeding 33

mm in standard length in old aquarium specimens

in N. marilynae and up to 35.3 mm in standard

length in old aquarium specimens of N. trijasciatus.

In live color pattern N. marilynae is most like N.

minimus but the males lack the highly modified

anal fin of the male N. minimus. In caudal-fin

modification but not in swimming position, N.

marilynae shows some similarities to two larger

species, N. unifasciatus (up to 38.8 mm in standard

length) and N. eques (up to 35.4 mm in standard

length). Both of these species are very different

in live as well as preserved color patterns from N.

marilynae.

ADDENDUM.—After this manuscript was set in

type, we received from Dr. Jamie Thomerson of

Southern Illinois University 190 mostly adult

specimens of Nannostomus marilynae, collected

2 April 1974 by J. E. Thomerson, Donald C. Hicks,

and Jesus Enrique Vaques from Cano Muco, about

15 kilometers west of Puerto Gaitan, toward Puerto

Carreno, state of Meta, Colombia. The Cano Muco

is a tributary of the Rio Vichada, which is itself a

tributary of the Orinoco River. These specimens

came from an area collected commonly by aquarium

fish exporters. This fact plus the detailed resemblance of these fishes to the aquarium specimens

of N. marilynae in Figures 29 and 30 make it

appear likely that these aquarium fishes came from

either this or some adjacent aquarium collecting

area in Colombia. The fact that N. marilynae occurs

in the Rio Negro, Brazil, and in at least one Colombian tributary of the Rfo Orinoco is not surprising. The headwaters of the tributaries of the

Orinoco closely approach the main courses of both

the Rio Infrida and Rio Guaviare to the south of

the Rio Vichada in Colombia. The Inirida and

Guaviare both drain into the Orinoco. Also the Rio

Orinoco and Rio Negro are connected by the Cano

Casiquiare. These closely lying rivers can be expected to have very similar faunas, and other collections recently acquired from Mr. Socolof of
NUMBER 186

Bratlenton, Florida, in the Rfo Vichada drainage

have borne this out. For example, one collection

from the Rfo Vichada drainage by Mr. Socolof has

produced the second known record of Brittanichthys axelrodi Gery, previously known only from

Praio Bulufu on the Rio Itu about 80 kilometers

up the Itu from where it meets with the Rio Negro

in Brazil. The locality is about 0°30' south latitude

and 63°30' west longitude, which places it in about

the center of the Rio Negro district. The cardinal

tetra, Cheirodon axelrodi Schultz, is also known

from these two areas in Brazil and Colombia. The

specimens in the Cano Muco collection of N. marilynae (not to be considered as paratypes) have been

distributed to the following museums: 125, FMNH

77981, SL 12.9-21.6 mm; 25, USNM 212161, SL

15.1-20.7 mm; 10, CAS 30868, SL 15.2-19.2 mm;

10, BM(NH), 1974.8.7.1-10, SL 15.8-20.2; 10,

ANSP 128438, SL 15.3-19.9; 10, ZMA 113.365, SL

14.8-19.8.

Nannostomus unifasciatus Steindachner

FIGURES 31, 32

DIAGNOSIS.—Primary horizontal stripe very well

developed, secondary and tertiary horizontal stripes

absent. Nocturnal oblique bands absent in preserved specimens; permanent blotches absent. Adipose fin present. Scales in a lateral series 28 to 30,

perforated scales in lateral line 2 to 5. Gill rakers

9+14 . Inner tooth row of dentary absent, teeth

in outer dentary row 6 to 7. Principal caudal-fin

rays 10/9 but those ending in dorsal lobe 8, those

ending in ventral lobe 11.

DISCUSSION.—We have here only the following

range extensions to add to the account of this

species given by Weitzman (1966).

Two, USNM 208258, SL 29.5-33.0 mm, BrazilBolivian border region, from the Rio Guapore

system somewhere between Guajara-Mirim and

the state border of Matto Grosso, 1970, Bernard

von Graeve. (These specimens have a welldeveloped caudal-fin ocellus.) 18, USNM 208259,

SL 24.9-31.5 mm, Brazil, Rio Purus at Boca de

Tapaua, H. Axelrod and H. Schultz.

Nannostomus eques Steindachner

FIGURES 33, 34

DIAGNOSIS.—Primary, secondary, and tertiary

horizontal stripes well developed, with an additional stripe on back above secondary stripe. Nocturnal oblique bands present; permanent blotches

absent. Adipose fin present or absent. Scales in a

lateral series 24 or 25. No perforated lateral-line

scales. Gill rakers 16 + 24. Teeth in inner dentary

row 12, teeth in outer dentary row 9. Principal

caudal-fin rays 10/9, but those ending in dorsal

lobe 8, while those ending in ventral lobe 11.

We have no information additional to that suplied by Weitzman (1966).

FIGURE 31.—Xannostomus unifasciatus, CAS(IUM) 11703, male, 33.4 mm SL, Wismar,

Demerara River, Guyana, 1908.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 32.—Xannostomus unifasciatus, male 35.6 mm SI., locality unknown, but specimen

compares well with specimens known to come from Guyana.

FK.IRK. 33.—Xannostomus eques, SU 50229 (now at CAS), male, 29.0 mm SL, rock pools below

rapids, Sao Gabriel, Rio Negro, Amazonas, Brazil, 1925.
NUMBER 186 35

FiGi'RE 34.—Xannostomus eques, female above, 33.4 mm SI.,

male below, 32.5 mm SL, locality unknown.

Sumario

Basado en material nuevamente acumulado,

este estudio es un tratamiento suplementario dc

una revista previa del ganero sudamericano Xannostomus por el aulor mayor. Once especies estan

reconocidas aqui, Ar. espei, X. diagrnmmus, A.

harrisoni, N. bechfordi, X. bifasciatus, X. minimus,

X. marginal us, X. Irifascialus, X. marilyntir

(nueva), N. unifasciatus y N. eques. Una clave

artificial a todas estas especies esta incluida. Colecciones adicionales de dos especies, N. bifascialus

y Ar. minimus, han permitido una descripcion

mas completa del aquella y reconocimiento de

las mas reciente que habia estado listada como

una sinonimo de N. beckfordi por Weit/man

(1966). Una nueva especie, N. marilynae, esta

descrita y ocurre en la cuenca amazona. Las corelaciones de todas las especies estan reexaminadas

y como la nueva material ha revelado condiciones

intermedias es los sujetos usados antes para separar

los generos nominales Nannoslomus y Poecilobrycon, este esta considerado como un sinonimo de

aquel. Las relaciones entre las especies no son

claras de las muestras poblaciones por lo general

disponibles para el estudio pero parece probable

que A7, espei en el total de sus sujetos es la especie

mas primitiva de Xannostomus y no esta relacionada solidamente ningunas otras especies. Xantwstomus marginatus y N. trifasciahts indudablemente se estan relacionadas solidamente pero no

se parecen mucho a otras especies o grupo de

especies. El mismo se puede decir de las relacionadas Ar

. eques y N. unifasciatus en sus relaciones

con otras especies. Las especies que se quedan,

parecen todas distinctas entre ellas y sus relaciones

posibles a otras especies en el genero, se quedan

obscuras.

Direcciones evolucionarias hacia la especializacion en el genero parecen ser el desarrollo de

rayas longitudinales y de rayas oblicas, la perdida

de la osificacion del canal sensorio en el segundo

hueso infraorbital, una elongacion y espesamiento

de las aletas rayas anales de los machos usadas

como una ayuda para fertilizar huevos, tal vez el

desarrollo de un ocelo en el lobulo dorsal de la

aleta caudal y de tubcrculos nupciales en la superficie ventral de la cabeza. Espesamiento de las

aletas rayas anales ocurre hasta cierto punto en

otros lebiasinidos pero esta llevado a extremo en

algunas especies de Xannostomus. Tuberculos

nupciales ocurren en otro genero lebiasinido,

Pyrrhulina, pero su gran concentracion en la

superfuie ventral de la cabeza en Ar. unifasciahis es

una especiali/acion no encontrada en Pyrrhulina.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Literature Cited

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92 + 332 pages, plates 634-650. Paris: Libraire de

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Eigenmann, C. H.

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1912. The Fresh-Water Fishes of British Guiana. Memoirs

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Fowler, H. W.

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Hocdeman, J. J.

1950. Rediagnosis of the Characid-Nannostomine Fish

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Hubbs, C. L.

1926. Studies of the Fishes of the Order Cyprinodontes,

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Meinken, H.

1951. Der Gold-Anomalus = Nannostomus anomalus

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Sterba, Giinther

1962. Freshwater Fishes of the World. 878 pages, 1193

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