The So-called Cheirodontin Fishes of

Central America with Descriptions of

Two New Species (Pisces: Characidae)

WILLIAM L. FINK

.

and

ANLEY H. WEITZMAN

m

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 172
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 17 2

The So-called Cheirodontin Fishes of

Central America with Descriptions of

Two New Species (Pisces: Characidae)

William L. Fink

and Stanley H. Weitzman

SMITHSONIAN INSTITUTION PRESS

City of Washington

1974
ABSTRACT

Fink, William L., and Stanley H. Weitzman. The So-called Cheirodontin Fishes

of Central America with Descriptions of Two New Species (Pisces: Characidae).

Smithsonian Contributions to Zoology, number 172, 46 pages, 26 figures, 15

tables, 1974.—The known species of Central American cheirodontin fishes,

characids with a single row of premaxillary teeth, are reviewed. A key to their

identification is provided. The review includes members of the genera Cheirodon,

Carlana, Phenagoniates, and Saccoderma. All species are described and illustrated

except for the Central American form of Saccoderma, which is known to us only

from juveniles and may belong to an undescribed species. Two new species of

Cheirodon are described: C. dialepturus from the Pacific drainages of west-central

and western Panama, from eastern Code Province west to the Rio Coto, Costa

Rica; and C. mitopterus from the Rio Code del None Basin, an Atlantic drainage of Central Panama.

Rhoadsia eigenmanni is placed in Carlana, the generic name available for

the preoccupied Carlia. The genera Odonlostilbe, Pseudocheirodon, and Compsura are synonomized with Cheirodon, based on reevaluation of the characters

formerly used to define these genera as well as a renewed evaluation of the

concept of the genus in characid fishes. Finally, some reasons are presented for

considering the Cheirodontinae polyphyletic.

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

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5054. SERIES COVER DESIGN:

The coral Monlastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Fink, William L.

The so-called cheirodontin fishes of Central America with descriptions of two new species

(Pisces: Characidae)

(Smithsonian contributions to zoology no. 172)

Supt. of Docs, no.: SI 1.27: 172.

1. Characidae. 2. Fishes—Central America. 3. Cheirodon dialepturus. 4. Cheirodon mitopterus.

I. Weitzman, Stanley H., joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 172.

QL1.S54 no. 172 [QL638.C5] 591'.08s [597'.'>2] 73-20338

For sale by the Superintendent of Documents, U.S. Government Printing Office

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Contents

Page

Introduction 1

Artificial Key to the Species of the So-called Cheirodontin Characids

of Central America 3

Cheirodon Girard 3

Cheirodon dialeptiirus, new species 5

Cheirodon mitopterus, new species 12

Cheirodon gorgonae Evermann and Goldsborough 14

Cheirodon affinis (Meek and Hildebrand) 19

Cheirodon terrabae (Bussing) 23

Carlana Strand 25

Carlana eigenmanni (Meek) 26

Phenagoniates Eigenmann and Wilson 30

Phenagoniates macrolepis (Meek and Hildebrand) 30

Saccodertna Schultz 33

Sumario 34

Tables . 36

Literature Cited 45

in

The So-called Cheirodontin Fishes of

Central America with Descriptions of

Two New Species (Pisces: Characidae)

William L. Fink

and Stanley H. Weitzman

Introduction

A survey of the characoid collections at the

National Museum of Natural History has produced many problems of identification. One of

these came to light when we examined a large

number of characid specimens from Panama, collected in 1961-1962, primarily by Dr. Horace Loftin.

A large percentage of the lots of so-called cheirodontins had been identified as Compsura gorgonae

(Evermann and Goldsborough), but close examination showed that many of these belonged to two

undescribed species. In an attempt to determine

the relationships of the new species, all known

Central American and many South American

cheirodontins were examined. To clear the way

for comparison with possible future discoveries in

Central America and to make information available for studies of South American cheirodontins,

we here present a review of all known Central

American species.

Our treatment of the Central American species

as cheirodontins is a matter of convenience and

tradition, and it is not to be interpreted that we

believe it is an expression of phylogenetic reality

William L. Fink and Stanley H. Weitzman: Division of Fishes,

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

Also, Fink: Department of Biological Sciences, George Washington University, Washington, D.C. 20006.

in the sense of Hennig (1966). In fact, we have

little confidence that the Cheirodontinae (American characids with a single row of premaxillary

teeth and lacking certain specializations of other

characoids; see Eigenmann, 1915) constitute a

group from a phylogenetic point of view. See

Bohlke (1952) for comments on the possible "polyphyletic" nature of the Cheirodontinae.

METHODS.—All measurements are straight-line

distances made with dial calipers, and all are in

standard length (SL) or percentages of standard

length unless otherwise indicated. Standard length

was measured from the anterior termination of

the upper jaw (tip of snout) to the center posterior

termination of the hypural fan (where the principal caudal-fin rays attach to the hypural bones).

Measurements referring to the origin of the fins

mean anterior (cephalad) termination of the fin

base unless otherwise stated. Fin length was measured from the fin origin to the distal tip of longest

ray. Head length equals the distance between the

tip of the snout and the posteriormost termination

of the bony opercle. (It is common to measure to

the posterior termination of fleshy opercular flap;

however, we find this flap to be damaged so frequently that a better measurement can be made

to the bony opercular border). Snout length extends from the tip of the snout to the anteriormost

bony edge of the orbit. Upper-jaw length was measured from the tip of the snout to the distalmost
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

termination of the maxillary bone. Eye diameter

is a longitudinal measurement between bony orbital margins. Distance from eye to dorsal-fin origin

is measured from posterior bony orbital margin.

Caudal-peduncle length was measured from posterior termination of the anal-fin base (at the insertion of last anal-fin ray) to the center of the

posterior termination of the hypural fan (end of

caudal peduncle; caudal-fin origin). Caudalpeduncle depth equals the least distance between

its dorsal surface and its ventral surface.

The greatest body depth extends from a point

along the dorsal profile to the ventral body surface

where the distance is greatest. The least bony interorbital width equals the least distance between

the dorsalmost bony edges of the orbits. Lateral

scales are counted as a longitudinal scale row along

the midline or along the perforated lateral-line

scales when these are present. Scales above the

lateral line are counted from the dorsal-fin origin

(not including the middorsal scale) to that scale

just dorsal to the midlateral (or perforated lateral)

scale row. Scales below the lateral line were counted

diagonally from the anal-fin origin (not including

scales of anal-fin sheath) to a scale just ventral

to the midlateral scale row. Scales around the

caudal peduncle were counted as the number of

scale rows intersecting a vertical line or ring around

the least depth of the caudal peduncle. The predorsal scales were counted beginning with the scale

at the posterior termination of supraoccipital spine

and continuing to the scale just anterior to the

dorsal-fin origin. In some cases the predorsal scales

were not counted due to a very irregular distribution. The last two dorsal-fin rays are not counted

as a single ray split to the base, but as two separate

rays. The last two anal-fin rays are counted as a

single ray split to the base; in many instances

these two rays are not completely split to the base.

(This method of counting dorsal- and anal-fin

rays reflects the actual anatomical relationships to

the pterygiophores.) Pectoral- and pelvic-fin rays

were all counted as separate rays. Gill rakers were

counted on both lower and upper limbs of the

first gill arch. Vertebral counts include the

Weberian apparatus as four elements.

The tooth figures were drawn from cleared and

stained specimens.

Specimens were selected to reflect the size range

exhibited at each collection locality.

In the descriptions, for each meristic character,

the range is given, followed by the mode; for each

morphometric character, the range is given. In

some descriptions, for morphometric characters,

juveniles and adults are separated by a semicolon.

In the "Material Examined" sections, an asterisk

placed before a collection number indicates that

the lot was used in the subsequent description.

Morphometric and meristic data are presented in

Tables 1-15; specimens included in tables are

noted following "Material Examined" under each

species. In tables, underscored numbers indicate

holotype meristics. Specimens damaged such that

counts or measurements for a given character were

not possible account for variation in number of

specimens in tables and descriptions. Following

the locality citation, Dr. Loftin's collection stations

(see Loftin, 1965) have been cross-referenced with

his station numbers (HL) and river basin numbers

(P or A) in parentheses. P stands for Pacific drainage and A for Atlantic drainage. IAH stands for

Inter American Highway.

Material was examined from collections in the

National Museum of Natural History (formerly in

United States National Museum: USNM), Washington, D.C., including material from the Battelle

Laboratory (BNW) through the Smithsonian

Oceanographic Sorting Center; Field Museum of

Natural History, Chicago (FMNH); California

Academy of Sciences, San Francisco (CAS); Stanford University (CAS[SU]), and Indiana University, (CAS[IUMJ), both collections now at California Academy of Sciences, San Francisco; Los

Angeles County Museum (LACM); Museum of

Comparative Zoology, Harvard University, Cambridge (MCZ); Academy of Natural Sciences of

Philadelphia, (ANSP); University of Michigan

Museum of Zoology (UMMZ); and Gulf Coast

Research Laboratory, Ocean Springs, Miss.

(GCRL). We wish to thank Reeve M. Bailey

(UMMZ), James E. Bohlke (ANSP), C. E. Dawson (GCRL), William Eschmeyer (CAS), Tyson

Roberts and Mrs. M. Dick (MCZ), and Loren P.

Woods (FMNH) for loan of specimens in their

care. C. E. Dawson loaned Los Angeles County

Museum material that was formerly deposited in

the University of Miami Ichthyological Museum

(UMIM) and is now at the Gulf Coast Research

Laboratory. We are grateful to J. D. McPhail and

R. L. Dressier for providing freshly preserved
NUMBER 172 3

specimens and color descriptions from life of two Central American characids. James E. Bohlke, J. D.

species and to W. A. Bussing for providing live McPhail, Reeve M. Bailey, Robert H. Gibbs, Jr.,

specimens of two species. George S. Myers pro- and W. A. Bussing read the manuscript and offered

vided a helpful and illuminating discussion of valuable suggestions.

Artificial Key to the Species of the So-called Cheirodontin

Characids of Central America

1. Anal fin with less than 40 branched rays. Dorsal-fin origin anterior to anal-fin origin. Teeth

usually with 5 or more cusps 2

Anal fin with more than 40 branched rays. Dorsal-fin origin posterior to anal-fin origin.

Teeth tricuspid Phenagoniates macrolepis (Meek and Hildebrand)

2. Anal fin with 15-23 branched rays. Scales below midlateral scale series 3-6 3

Anal fin with 28-30 branched rays. Scale rows below midlateral scale series 7-8

Carlana eigenmanni (Meek)

3. Modified caudal scales when present not elongate, not passing far onto lower caudal-fin lobe.

Dentary teeth with 5-10 narrow cusps (Figure 2) 4

Modified caudal scales elongate, passing well into lower caudal-fin lobe (Figure 26). Dentary

teeth with 6-8 broad triangular cusps forming slightly rounded cutting edge (Figure 25)

Saccoderma species

4. Premaxillary teeth approximately symmetrical, anterior or median and posterior or lateral

cutting edges about equidistant from tooth base (Figure 2). Dentary teeth with 5-10 cusps,

with middle cusp of tooth largest, forming a rounded cutting edge. Caudal peduncle or

caudal fin with modified scales1 5

Premaxillary teeth asymmetrical, with anterior or median cutting edge beginning closer to

tooth base than posterior cutting edge (median one or two teeth may be symmetrical) (see

Figure 13). Dentary teeth with 6-8 cusps, lacking an elongate middle cusp, forming an

even cutting edge. No modified caudal scales 7

5. Modified caudal scales not forming "pouches," consisting of two irregular ventral rows of

small terminal peduncle scales and several scales extending slightly onto middle caudal-fin

rays 6

Modified caudal scales forming three obvious "pouches" (Figure 10)

Cheirodon gorgonae Evermann and Goldsborough

6. Recurved bony hooks present on upper (dorsal) fin rays of lower (ventral) caudal-fin lobe

of males. Branched pectoral-fin rays 10-11, usually 10. Unbranched pelvic ray not filamentous in either sex Cheirodon dialepturus, new species

No recurved bony hooks on lower caudal-fin lobe. Branched pectoral-fin rays 11-12, usually

11. Unbranched pelvic-fin ray filamentous in males Cheirodon mitopterus, new species

7. Branched anal-fin rays 17-22 (x = 19.6). Dentary with about 7 broad multicuspid and 2-3

smaller conical teeth. Gill rakers 17-20 (X = 18.0). Lateral line usually incomplete

Cheirodon affinis (Meek and Hildebrand)

Branched anal-fin rays 21-23 (x = 22.1). Dentary with about 8-9 broad multicuspid teeth

and usually 2-3 smaller conical teeth. Gill rakers 19-21 (x = 20.4). Lateral line usually

complete Cheirodon terrabae (Bussing)

1

 Modified caudal peduncle scales may be difficult to see in C. dialepturus and C. mitopterus;

the scales are easily damaged.

Cheirodon Girard Pseudocheirodon Meek and Hildebrand, 1916:275 [type-species

Pseudocheirodon affinis Meek and Hildebrand, 1916, by

Cheirodon Girard, 1854:199 [type-species Cheirodon piscicuhis original designation and monotypy].

Girard, 1854, by monotypy].

Odontostilbe Cope, 1870:566 [type-species Odontostilbe fugi- -phe generic referral below of five species of Centiva Cope, 1870, by monotypy]. tra l America n " che irodontins" to Cheirodon needs

Compsura Eigenmann, 1915:60 [type-specics Compsura hete- MIL

rura Eigenmann, 1915, by original designation and explanation. The two new species described here

monotypy]. as new, plus reexamination of other Central
American "cheirodontins," has forced a reconsideration of the generic validity of Odontostilbe,

Compsura, and Pseudocheirodon.

One species herein described as new, Cheirodon

mitopterus, could be referred to the genus Odontostilbe Cope (1870) because it has a complete

lateral line. Cope described Odontostilbe for O.

fugitiva Cope (1870) because at that time it was

the only known characid with a single row of premaxillary teeth and a complete lateral line. G£ry

(1972:71) has most recently denned Odontostilbe

and implied in his discussion that it differs from

Cheirodon only in the possession of a complete

lateral line. We can no longer accept the possession

of a complete versus incomplete lateral line as a

valid generic character separating Cheirodon and

Odontostilbe because some specimens of two of

the species of Cheirodon treated here, C. dialepturus (new species) and C. affinis (Meek and Hildebrand), have complete and others have incomplete

lateral lines. Cheirodon dialepturus usually tends

to have an incomplete lateral line, but some population samples usually have a complete lateral line.

Specimens of most population samples of C. affinis

tend to have an incomplete lateral line, but some

samples have all or nearly all of the specimens

with a full row of 33 perforated lateral-line scales.

In view of the demonstrated lability of the lateralline length, we see no reason to recognize the genus

Odontostilbe, and therefore consider it a synonym

of Cheirodon. See below under Cheirodon dialepturus, C. affinis, and C. terrabae (Bussing), for a

more complete discussion of lateral-line variation.

Compsura has been defined as a "cheirodontin"

with modified scales associated with one or more

well-developed pouches on the lower caudal-fin

lobe. In addition, the known species have teeth

with symmetrically arranged cusps, each tooth

having an enlarged median cusp. Retrorse hooks

are present on the anal- and pelvic-fin rays in males

and the lower (ventral) caudal-fin rays are without hooks. This definition fits both C. heterura

Eigenmann, the type-species of the genus, from

eastern Brazil (Rio San Francisco and Rio Itapicuru) and the fish heretofore known as C. gorgonae

(Evermann and Goldsborough), with a limited distribution in central Panama (see below for collection records). The wide geographical separation

of these two species, nearly 3,000 miles, strains

credulity that they must be one another's closest

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

relatives as indicated by their common generic

status.

If the characters used to define Compsura are

labile ones appearing more or less at random

among small characids, convergence of characters

would seem a better hypothesis than one of close

relationship to explain the similarities of these

two species separated by such a wide geographical

distance. The only character in the definition

above that has not been demonstrated to be labile

at the species level is the caudal-fin squamation.

Even here, in view of the possible independent

appearance and perhaps even lability of the caudal

region in several of the so-called "glandulocaudins"

(Weitzman and Thomerson, 1970) and the fact

that subtle modifications in scale structure occur

in other species of Cheirodon (see accounts of C.

mitopterus and C. dialepturus below), we do not

believe that the caudal glands and associated scales

in characoids are necessarily useful in determining

generic relationships.

Accordingly, the holotype (FMNH 57825) and

several other specimens of Compsura heterura

(FMNH 57827, 57828) have been examined and

compared with C. gorgonae to determine whether

the two are truly congeneric and the caudal scales

are similar enough that convergence would seem

unlikely. None of the C. heterura specimens available have all the caudal scales present (some have

been dried in the past and are quite damaged),

but an approximate reconstruction of the caudalscale configuration was made (Figure 11). The figure of the caudal fin in Eigenmann (1915:61)

apparently is not accurate in that some of the

scales are misrepresented or missing. The modified

caudal scales are not identically arranged in the

two species under consideration as may be seen

by comparing Figures 10 and 11. In C. heterura the

scales are more elongate and associated with a

single large pouch on the ventral part of the caudal

peduncle. A modified scale covers a large area of

the base of the dorsal caudal-fine lobe. In C. gorgonae there is a series of 3 layers of scales, each

associated with its own pouch, all in relation to

the ventral part of the caudal peduncle. There is

little modification in the scales above the midline

in C. gorgonae. No attempt was made by histological methods to determine whether glandular tissue

is associated with the pouches in either species,

but glandular tissue is not obvious to gross inspec-
NUMBER 172

tion. Eigenmann (1915) indicated that only males

of C. heterura possess modified scales, whereas we

found these scales in both sexes of C. gorgonae.

All specimens of C. heterura examined by us were

males, except for 2 small juveniles, 1 of which had

enlarged scales on the caudal peduncle.

Thus the principal character that separates these

two fishes, C. heterura and C. gorgonae, from members of the genus Cheirodon appears to be very

different in each species and likely developed independently in both. The remaining characters

supplementally defining Compsura appear to us

to be labile characters appearing here and there

in the species of small characids, and although

they have, in various combinations, been used to

help define genera, their use in defining relationships is open to question. If one wishes to refer

C. heterura to a genus separate from Cheirodon

because of its caudal structure, the same should

be done for C. gorgonae, and since the caudal

structure is different in the two species, they would

have to be placed in separate genera. In view of

the difficulty of using caudal structures as generic

characters within the so-called glandulocaudins,

and the subtle but certainly evident caudal-fin

variation and modification in other members of the

genus Cheirodon, we are here considering both

C. heterura and C. gorgonae as members of the

genus Cheirodon pending a full investigation of

caudal scale and pouch structure in characid fishes.

Two of the species here referred to Cheirodon—

C. affinis and C. terrabae—could be placed in the

genus Pseudocheirodon Meek and Hildebrand (C.

affinis is the type-species of the genus). Bussing

(1967) and Lopez (1972) accepted Pseudocheirodon

as valid because of an even cutting edge of the

dentary teeth, lack of fin hooks, and low number

of "interhemals" (ventral procurrent caudal-fin

rays) as compared with the rounded cutting edge

of the dentary teeth, presence of anal-fin hooks,

and enlarged, numerous (up to 20) ventral procurrent caudal-fin rays in Cheirodon. These characters as treated by Bussing need some comment.

We have found anal-fin hooks in C. affinis and C.

terrabae. As pointed out by Bohlke (1954), the

number and size of ventral procurrent caudal-fin

rays vary greatly in Cheirodon, and we believe it

to be a labile character that should not be used

as a generic character for this group of fishes. Bussing also felt hesitant about using ventral procurrent caudal-fin rays as a generic character.

The remaining character, an even cutting edge

in each dentary tooth in Pseudocheirodon, versus

an enlarged median or central cusp and a rounded

cutting edge in each dentary tooth in all other

species of Cheirodon does not seem valid. We find

that some specimens of at least C. affinis (the typespecies of Pseudocheirodon) have somewhat enlarged median central cusps on the dentary teeth,

approaching some undescribed species of Cheirodon from northern South America, which tend to

have a relatively small central median cusp. The

character as recorded for Pseudocheirodon is not

sharply different from that found in some species

of Cheirodon, and all the characters so far noted

that might separate Pseudocheirodon from Cheirodon appear to grade between the two nominal

genera.

In our key we have separated C. affinis and C.

terrabae from other Central American species of

Cheirodon on the basis of an asymmetrical cutting

edge of the premaxillary teeth. This character is

useful in the key on a local geographical basis;

however, since several other species of Cheirodon

from elsewhere have asymmetrical teeth, some

subtly so, some obviously so, we believe that this

character is labile and useless at the generic level.

Cheirodon dialepturus, new species

FIGURES 1-5

Compsura gorgonae (not Evermann and Goldsborough).—Nelson, K., 1964:62 [in part, listed], 71, fig. 3a [caudal morphology figured], 74 [in part, identity of specimens discussed], 129 [in part, discussion of possible relationship

with the Glandulocaudini].—Bussing, 1967:211 [listed],

214 [in part, compared with Pseudocheirodon], 241, fig. 1

[photograph].—L6pez, 1972:93-129 [in part, compared with

C. affinis and C. terrabae; variation of several characters;

distribution], 121, fig. 5 [photograph].

MATERIAL EXAMINED

(All specimens from Panama except as noted)

•USNM 208524, holotype, a male 28.0 mm: Veraguas

Province, Rio San Pedro basin, creek at bridge 12 mi W

of Santiago on Road to Sona, H Loftin, E. Tyson, R.

Yerger, 28 Jan. 1962. (HL-105; P16-9).

•USNM 208523, paratypes, with same data as holotype; 144

specimens 20.1-30.0 mm (3 cleared and stained).

•ZMA 112.473, paratypes, with same data as holotype; 2

specimens 26.3-26.5 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 1 Cheirodon dialepturus, new species, holotype, USNM 208524, male, 28.0 mm; Panama,

Veraguas Province, Rio San Pedro basin, creek at bridge 12 miles west of Santiago on road to

Sona, collected 28 January 1962 by H. Loftin, E. Tyson, and R. Ycrger.

•BMNH 1973.2.161; 3-4, paratypes, with same data as holotype; 2 specimens 25.8-28.8 mm.

•FMNH 71702, paratypes, with same data as holotype; 2

specimens 27.4-29.6 mm.

•CAS 16060, paratypes, with same data as holotype; 2 specimens 28.3-30.0 mm.

•MCZ 49067, paratypes, with same data as holotype; 2 specimens 26.6-28.3 mm.

•ANSP 121989, paratypes, with same data as holotype; 2

specimens 28.4-29.3 mm.

•USNM 208525, Veraguas Province, Rio San Pedro basin,

Rio Cobre at bridge on new IAH section from Santiago,

H. Loftin, E. Tyson, R. Condon, 4 Jan. 1962; 16 specimens

24.9-30.7 mm (HL-92; P17-3).

•USNM 208522, Chiriqui Province, creek 5 mi W of David

on IAH, H. Loftin and E. Tyson, 2 Dec. 1961; 94 specimens

18.8-33.0 mm (2 specimens cleared and stained) (HL-72;

P21-10).

USNM 208552, Code Province, Rio Grande basin, creek of

Rio Cocle about 5 mi N of Penonome on road to La

Pintada, H. Loftin, 23 Mar. 1962; 17 specimens 14.8-29.7

mm (HL-134; P10-1).

USNM 208531, Cocle Province, Rio Hato basin, creek, about

3 mi \V of Rio Hato at bridge on IAH, H. Loftin and E.

Tyson, 14 Oct. 1961; 7 specimens 16.5-27.0 mm (HL-16;

P9-18).

USNM 208528, Cocle Province, Rio Grande basin, creek at

bridge on IAH about 5 mi E of Nata, H. Loftin and E.

Tyson, 15 Oct. 1961; 18 specimens 19.7-26.3 mm (HL-22;

PI 0-4).

USNM 208538, Cocle Province, creek on IAH about 3 mi E

of Davisa, H. Loftin and E. Tyson, 15 Oct. 1961; 8 specimens 16.0-18.9 mm (HL-25; PI 1-3).

USNM 208509, Veraguas Province, Rio Santa Maria basin,

creek 1 mi S of Rio Santa Maria bridge on San Francisco

road, H. Loftin, 14 Jan. 1962; 20 specimens 17.2-22.3 mm

(1 cleared and stained) (HL-97; PI 1-13).

USNM 208536, Veraguas Province, Rio San Pedro basin,

creek at bridge about 20 mi W of Santiago on Sona road,

H. Loftin and E. Tyson, 28 Oct. 1961; 56 specimens 16.8-

27.3 mm (HL^O; P16-12).

USNM 208526, Veraguas Province, Rio San Pedro basin, creek

about 21 mi W of Santiago on Sona road, H. Loftin and

E. Tyson, 28 Oct. 1961; 3 specimens 26.4-27-3 mm (HL-41;

P16-13).

USNM 208512, Veraguas Province, Rio San Pablo basin,

creek 2 mi W of Rio San Pablo bridge on new IAH section

from Santiago, H. Loftin, E. Tyson, R. Condon, 4 Jan.

1962; 17 specimens 26.4-28.9 mm (2 cleared and stained)

(HL-91; P17-1).

USNM 208532, Chiriqui Province, backwater of Rio San

Felix by bridge on IAH (new section), H. Loftin and E.

Tyson, 18 Nov. 1961; 61 specimens 17.2-26.2 mm (HL-63;

P20-7).

USNM 208537, Chiriqui Province, creek on IAH (new section)

less than one-fourth mile from San Felix bridge, H. Loftin

and E. Tyson, 18 Nov. 1961; 93 specimens 22.3-31.3 mm

(HL-62; P20-8).

USNM 208527, Chiriqui Province, Rio Chorcha at bridge on

IAH E of town of Chiriqui, H. Loftin and E. Tyson, 1 Dec.

1961; 95 specimens 27.9-32.2 mm (HL-71; P2O-17).

USNM 208535, Chiriqui Province, Rio Esti, about 1 mi N of

Gualaca, H. Loftin and E. Tyson, 16 Dec. 1961; 10 specimens 23.1-34.3 mm (HL-80; P21-1).

USNM 208510, Chiriqui Province, Creek 15 mi W of Concepcion on IAH, H. Loftin and E. Tyson, 2 Dec. 1961; 17

specimens 25.0-30.0 mm (2 cleared and stained) (HL-77;

P22-1).
NUMBER 172

USNM 208508, Chiriqui Province, 1.45 mi E of Callc Segunde

este, La Concepcion on I AH, R. H. Goodyear, 16 Apr.

1969; 17 specimens 27.1-30.1 mm (2 cleared and stained).

USNM 208511, Panama Province, Rio Corona at bridge on

IAH 3 mi E of San Carlos. H. Loftin and E. Hislop, 11

Mar. 1962; 14 specimens 27.8-30.0 mm (2 cleared and

stained) (HL-127; P9-10).

USNM 208530, Panama Province, Rio Las Lajas at bridge on

IAH E of San Carlos, H. Loftin, 24 Mar. 1962; 2 specimens

30.4-31.8 mm (HL-136; P9-6).

USNM 208533, Comoro del Baru Province, large irrigation

canal by road 8 mi N of Puerto Armuelles, H. Loftin and

E. Tyson, 15 Apr. 1962; 5 specimens 28.1-29.7 mm (HL148; P23-1).

USNM 208529, Los Santos Province, Rio La Villa basin, Rio

Tebarico, about 3 mi W of Llano de Peidra, H. Loftin

and E. Tyson, 30 Sept. 1961; 63 specimens 22.0-29.5 mm

(HL-08; PI2-1).

USNM 208534, Herrera Province, creek about 4 mi up Pese

road from junction with Chitre-Divisa road, H. Loftin

and E. Tyson, 21 Oct. 1961; 113 specimens 15.8-29.1 mm

(HL-32; P12-3).

USNM 210990, Costa Rica; Puntarenas Province, 7 km SE

of Villa Neily on IAH, M., E., and W. Bussing, 28 Feb.

1973; 4 specimens 27.9-32.6 mm.

Specimens included in description and tables

include holotype, 6 male and 4 female paratypes,

6 males and 4 females each from USNM 208525

and USNM 208522.

DESCRIPTION.—Standard length of examined

specimens 25.4-33.3 mm. Body elongate, compressed

POSTERIOR (LATERAL)

CUTTING EDGE

ANTERIOR (MEDIAN)

CUTTING EDGE

FIGURE 2.—Dentition of Cheirodon dialepturus, USNM 208522,

female, 31.5 mm; Panama, Chiriqui Province, creek 5 miles

west of David on IAH, collected 2 December 1961 by H.

Loftin and E. Tyson.

laterally; greatest body depth 32.2-38.3. Predorsal

body profile somewhat convex with slight concavity at nape; concavity deepest at posterior termination of supraoccipital spine. Body profile

between posterior dorsal-fin base termination and

anterior base of adipose fin nearly straight to slightly convex, often with a slight dip at adipose fin

base. Posterior to adipose fin, body profile continuous as a straight line or slight ventrally directed

slope to procurrent caudal rays. Distance from eye

to dorsal-fin origin 38.0-41.2; distance from dorsalfin origin to end of caudal peduncle 51.3-54.7.

Ventral body profile usually gently rounded from

jaws to anus; steepest inclination ventral to jaws.

Ventral body profile protrudes ventrally its greatest

distance just anterior to pelvic-fin origin. Body

profile along anal-fin base straight or slightly concave; between posterior anal-fin termination and

procurrent caudal rays, body profile concave. In

many males ventral procurrent caudal rays slightly

longer than dorsal procurrent rays. Caudalpeduncle depth 12.1-15.0; peduncle length 13.9-

16.5.

Head Length 23.3-25.3. Eye diameter 8.6-10.5.

Snout length 4.7-6.1. Least bony interorbital width

7.0-8.0. Maxillary relatively long, sloping ventrally

and posteriorly, forming an angle of 50-60 degrees

to longitudinal body axis; upper-jaw length 7.1-

9.4. All teeth symmetrical, with 6-7 cusps (except

the small posterior tricuspid or conical dentary

teeth), with median cusps longest and at least 1

enlarged median cusp; all teeth in a single series

(see Figure 2). Maxillary with 2-3, usually 3, teeth

(very rarely with 4 teeth), premaxillary with 5

teeth (2 specimens with 6 on one side). Dentary

with 9-11 teeth, broader than those of maxillaries,

arching anteriorly and becoming smaller toward

rictus, row ending in a small conical tooth. No

teeth present on vomer, palatines, or pterygoids.

Fontanel moderate, that part anterior to epiphyseal bar about as long as width of fontanel

posterior to bar. Gill rakers moderate, 18-21. Circumorbital bones well ossified, infraorbital 3 wide,

contacting preopercle ventrally, and with a narrow

naked posterior area. Infraorbital 2 not touching

preopercle as indicated by Lopez (1972) for "gorgonae" (see below concerning identity of her specimens).

Scales moderately large, cycloid, with concentric

circuli, and about 4-6 radii on exposed posterior
8

field. Lateral line variable, with as few as 7 perforated scales to as many as 34 in some specimens

(holotype with 13 perforated scales). When complete, lateral line with a ventral curve on side of

body to below adipose fin, then continuing as a

straight line to caudal-fin base just ventral to midline. Lateral scales 33-34, usually 33 (often irregular and difficult to count with accuracy); scales

above lateral line 6 (some populations examined

with 5 scales above lateral line); scales below lateral

line 4 in all specimens. Predorsal scales 10-12,

usually 11. Scale sheath at base of anal fin of about

5-6 scales. Axillary scale present dorsal to pelvicfin insertion. Posterior 2 irregular layers of caudalpeduncle scales (border scales) smaller than those

just anterior (Figure 3). Each of 2 or 3 posteriormost scales at midline longer than deep, these elonSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

gate scales overlapping smaller adjacent scales. All

caudal-peduncle scales combined protruding slightly

out over caudal fin and attaching to fin membrane. Some specimens with a complete lateral

line, with an extension of lateral line onto caudal

fin. Variation in exact scale arrangement present,

but all specimens have elongate scales at midline.

Dorsal fin with 2 unbranched rays and 8-9, usually 9, branched rays. Dorsal-fin origin anterior to

anal-fin origin, posterior to pelvic-fin origin, nearer

eye than caudal-fin base. Distance from tip of snout

to dorsal-fin origin 50.5-55.2. Second or third ray

of dorsal fin longest with posterior rays shorter,

forming a slightly rounded posterior margin to

fin; length of longest ray 27.3-33.8 (holotype damaged).

Anal fin with 4 unbranched rays and 16—19,

Fir.t'RE 3.—Caudal fin and adjacent scalatiun of Cheirodon dialepturus, USNM 208:">2, male, 26.5

mm; Panama, Rio Grande basin, creek of Rio Code about 5 miles north of Penonomc on road

to La Pintada, collected 23 March 1962 by H. Loftin.
NUMBER 172

usually 17 or 18, branched rays. First unbranched

ray not always visible except in radiographs. Origin

of anal fin, posterior to midpoint of standard

length, 62.1-66.4. Fourth through eighth or ninth

anal rays longer, with successive posterior rays

shorter, forming an abruptly protruding fin margin

anteriorly and a straight margin posteriorly; this

character more pronounced in females than in

males. Dorsally recurved anal-fin hooks present in

males only; these occur on posterior unbranched

ray and on posterior branches of first through

fourteenth (1 specimen), fifteenth (8 specimens,

including holotype), sixteenth (5 specimens), or

seventeenth (2 specimens) branched rays; only 1

hook or bilateral pair of hooks per bony ray segment; fin often fleshy around hooks. Posterior 8 or

9 branched anal-fin rays often bend slightly anteriad. Extent of area covered by anal-fin hooks

and bending of posterior rays varies from population to pojjulation.

Pectoral fin with 1 unbranched ray and 9-11,

usually 10, branched rays. Pectoral fins reach just

anterior to or just to pelvic-fin origin. Distance

from tip of snout to dorsal end of pectoral-fin base

23.4-26.8 and length of pectoral fin from base to

tip of longest ray 20.8-26.2.

Pelvic fin with i,7 rays in all specimens, distal

tip reaching from just anterior to just posterior

to anal-fin origin. Distance from tip of snout to

pelvic-fin origin 43.7-48.8; pelvic-fin length 17.9—

24.2. In males, unbranched ray without hooks, first

through sixth (2 specimens) or seventh (13 specimens including holotype) branched rays with

antrorse hooks, usually on the ventral surface of

rays. Only 1 hook present per bony ray segment.

Caudal fin with 10/9 principal caudal rays in

all specimens; fin forked, not split to base. Lower

lobe largest in males, with recurved hooks present

on dorsal edge (occasionally ventral edge) of

branches of rays 2-5 or 2-6, counting the most

dorsal ray of lower lobe as number 1. These hooks

often paired, similar to anal-fin hooks, with each

hook of pair facing different side of fin; only 1

hook or hook pair per bony ray segment. Fin rays

usually curved dorsally when hooks present. Ventral procurrent rays of males often somewhat

longer than dorsal procurrent rays. In females,

caudal fin symmetrical, without hooks and elongate

ventral procurrent rays.

Precaudal vertebrae 15-16. Total vertebrae 32-

34, usually 33.

Color in alcohol: Ground color pale brown;

nape dark brown. Humeral spot variable, virtually

absent in some populations, rather large in others;

FIGURE 4.—Cheirodon dialepturus, new species, USNM 210990, male, 32.5 mm; Costa Rica,

Puntarenas Province, 7 kilometers southeast of Villa Neily on I AH, collected 28 February 1973

by Myrna, Eric, and William Bussing; specimen photographed in life.
10

when present, centered at "pseudotympanum."

Small melanojihores present on scale pockets on

sides of fish, more numerous dorsally, forming

reticulate pattern above midline and following

myomere junctions below midline above anal fin.

Some specimens with a broad dark stripe along

midline, terminating in dark caudal spot, in others

only caudal spot present. Small melanophores

usually present along fin rays of all fins, more

numerous distally on dorsal, caudal, and anal fins.

Color hi life: The following description has

been supplied to us by Dr. J. D. McPhail from

specimens captured in the Rio Zarati, on road

between Penonome and Tambo, Code Province:

Back bronze shading to green. Scales of back outlined with black pigment and very conspicuous.

Belly silver white, sides dorsal to anal fin translucent. Burnished silver lateral stripe ending at

dark spot on caudal peduncle. Very conspicuous

yellow pigment (shading to red in some males)

above and below caudal spot. Dorsal, anal, and

pelvic fins red with white tips. Caudal fin red,

shading off to transparency toward fin margin.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Range: Cheirodon dialepturus is found in Pacific drainages throughout western and west-central

Panama from eastern Code Province, west at least

to Puntarenas Province of Costa Rica (see "Material Examined"). Bussing (1967) listed C. gorgonae from the Rio Coto drainage in eastern Costa

Rica; Lopez (1972) approximated C. gorgonae's

range to be from the Rio Coto, throughout southern Panama, to areas of Colombia adjacent to

Panama (no specimens were listed from areas east

of the Canal Zone). We believe, as discussed below,

that Lopez and Bussing were referring, for the

most part, to C. dialepturus.

COMPARISONS.—Cheirodon dialepturus is a distinctive species that can be separated from all

other known Central American cheirodontins by

the combination of the presence of antrorse hooks

on the ventral caudal-fin lobe of males, and the

modified scales on the caudal peduncle as shown

in Figure 3.

Cheirodon dialepturus has jaw teeth similar to

those of C. gorgonae but lacks the modified scale

"pouches" of the latter. Also, the two species have

FIGURE 5—Cheirodon dialepturus, new species, USNM 210990, female, 32.6 mm; Costa Rica,

Puntarenas Province, 7 kilometers southeast of Villa Ncily on IAH, collected 28 February 1973

by Myrna, Eric, and William Bussing; specimen photographed in life.
NUMBER 172

a different body shape (compare Figures 1 and 8).

Cheirodon dialeplurus resembles C. mitoptcrus in

its possession of modified caudal-peduncle scales;

however, other characters including live color pattern, fewer vertebrae (32-34, usually 33, in C.

dialepturus, vs. 34-36, usually 35, in C. miiopterus),

fewer pectoral rays (9-11, usually 10, in C. dialepturus, vs. 11-12, usually 12, in C. mitopterus),

fewer tooth cusps (6-7 in C. dialepturus, vs. 7-10

in C. mitopterus), and presence of caudal-fin hooks

(lacking in C. mitopterus) clearly set the two

species apart. Furthermore, there are definite behavioral differences between the two forms (McPhail, personal communication). Cheirodon dialepturus shares the characters of caudal-fin hooks with

some species of Saccoderma (discussed in the account of that genus) but is clearly different from

those species in tooth structure and in the shape

of the latter's modified caudal-fin scales. Cheirodon

dialepturus is quite distinct from C. affinis and

C. terrabae in having caudal-fin hooks, a different

tooth structure, and the modified caudal scales.

There has been some confusion regarding C.

dialepturus in the past. Loftin's collections of the

species in the National Museum of Natural History

are mostly labeled "Compsura gorgonae" or "Pseudocheirodon affinis." Loftin (1965) states that his

cheirodontins were examined and identified by W.

A. Bussing. Therefore, we assume that much of

Loftin's and Bussing's (1967) "Compsura gorgonae"

material may be C. dialepturus. This is substantiated by the fact that Bussing refers to the hooks on

the lower caudal lobe of his specimens. We also

note that Gery (1965) apparently confused these

two species when he referred to caudal-fin hooks of

Compsura gorgonae. Lopez (1972) also confused C.

dialepturus and C. gorgonae; she referred to caudal

hooks and pouches in C. gorgonae and apparently

thought that their presence or absence was due to

intraspecific variation.

VARIATION.—The extensive collections of C.

dialepturus listed above in "Materials Examined"

have been used to determine the geographical range

of the species, individual and population differences, and seasonal morphological changes. Variation in C. dialepturus is most evident in

pigmentation, tooth structure, and number of

lateral-line pores. As mentioned in the color description of alcohol specimens, some populations

have a broad lateral stripe and a rather pronounced

11

humeral spot, while others lack both but have a

caudal spot. Within a population some individuals

will have one or the other or a particular combination of these characters more or less pronounced

than usual for that population. The slight variation in tooth structure occurs mainly in the massiveness or length of the tooth cusps. This tooth

variation, for the most part, seems to follow no

particular geographical or ecological pattern, except that those specimens from western Panama

seem to have slightly broader teeth with smaller

cusps. The most obvious morphological variation

is in the number of lateral-line pores. Specimens

from central Panama have incomplete lateral lines

(extending posteriorly to a point well anterior to

a vertical or just anterior to a vertical from the

dorsal-fin origin). Population samples of C. dialepturus from extreme western Panama tend to have

some or all individuals with complete or nearly

complete lateral lines. However, some collections

from this area have all specimens with incomplete

lateral lines similar to those in fishes from central

or west-central Panama. Contrary to Lopez (1972),

we could find no ecological factors among the field

data recorded by Loftin, such as vegetation, elevation, water flow, turbidity, or depth, which could

be associated with the number of lateral-line pores.

We have found no seasonal variation in C. dialepturus regarding coloration or massiveness and

length of anal, pelvic, or caudal-fin hooks.

Lopez (1972) stated that there was geographical

variation in length of the maxillary bone in what

now has been found to be mixed samples of C.

gorgonae, C. dialepturus, and C. mitopterus. We

found no significant correlation with upper-jaw

length and geographical distribution in these

species. There is, as would be expected, a slight

size increase in total upper-jaw length with increase

in body length and our data suggest that this is

an allometric increase. The small size and difficulty

of measuring the length of the maxillary bone as

outlined by Lopez (1972) would require cleared

and stained specimens and the use of an ocular

micrometer to obtain useful measurements. We

believe extreme care should be used in measuring

such small structures.

ETYMOLOGY.—From the Greek dialeptos (distinguishable) and oura (tail), referring to the

caudal hooks and peduncle scalation.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Cheirodon mitopterus, new species

FIGURES 6, 7

Compsura gorgonae (not Evermann and Goldsborough).—

L6pez, 1972:101 [listed from Rio Code del Norte, HL-172],

106 [discussion of lateral-line length].

MATERIAL EXAMINED

(All specimens from Panama)

•USNM 208539, holotype, a male 35.0 mm: Code Province,

Rio Code del Norte basin, Rio Tucue at junction of river

and road between Tucue and Tambo, coll. J. D. McPhail,

28 Aug. 1972.

•USNM 208513, paratypes, with same data as holotype; 7

specimens 28.5-36.8 mm (1 cleared and stained).

•ANSP 121987, paratype, with same data as holotype; 1

specimen 30.9 mm.

•BMNH 1973.2.16;1, paratype, with same data as holotype;

1 specimen 30.4 mm.

USNM 208540, Code Province, Rio Code del Norte basin, Rio

Tucue, near village of Tucue in headwaters of Rio Code

del Norte, H. Loftin and W. Kosan, 15 Sept. 1962; 12

specimens 25.7-33.4 mm (HL-171; A5-1).

USNM 208514, Code Province, Rio Code del Norte basin,

Rio Tambo, near Tambo and Toabre, in headwaters of

Rio Code del Norte, H. Loftin and W. Kupfer, 15 Sept.

1962; 14 specimens 24.6-33.1 mm (2 cleared and stained)

(HL-172; A5-2).

USNM 208541, Code Province, Rio Code del Norte basin,

stream near Toabre road (either the Rio Toabre or a

small tributary of it) near Tambo, R. Dressier, Sept. 1972;

12 specimens 28.4-35.1 mm.

Tables include 10 specimens used in the description and 10 specimens each from USNM 208540

and USNM 208514.

DESCRIPTION.—Standard length of examined

specimens 24.9-36.8 mm. Body elongate, compressed laterally; greatest body depth 23.6-38.3.

Predorsal body profile somewhat convex, with a

slight concavity at nape. Body profile nearly straight

to slightly convex between posterior dorsal-fin base

termination and anterior base of adipose fin. Posterior to adipose fin, body profile continuous as a

straight line to procurrent caudal-fin rays. Distance

from eye to dorsal-fin origin 29.8-49.7; distance

from dorsal-fin origin to caudal-fin origin 44.0-74.3.

Ventral body profile very gently rounded from jaws

to anus, then with a slight ventral slope to analfin origin; steepest inclination of ventral profile

ventral to jaws. Ventral body profile protrudes

ventrally its greatest distance just anterior to pelvicfin origin. Body profile along anal-fin base straight

or very slightly convex; between posterior anal-fin

termination and procurrent caudal rays body profile slightly concave. Caudal peduncle depth 8.5-

14.7; peduncle length 13.1-21.0.

Head length 18.5-28.7. Eye diameter 7.9-12.9.

Snout length 4.3-7.4. Least bony interorbital width

5.7-9.2. Maxillary moderate, sloping ventrally and

posteriorly, forming an angle of about 50 degrees

to longitudinal body axis; upper-jaw length 6.2-

9.9. All teeth symmetrical, with 7-10 cusps (except

the small posterior dentary teeth discussed below),

with median cusps longest and at least 1 enlarged

^n-O^:

FIGURE 6.—Cheirodon mitopterus, new species, holotype, USNM 208539, male, 35.0 mm; Panama,

Code Province, Rio Code del Norte basin, Rio Tucue at junction of river and road between

Tucue and Tambo, collected 28 August 1972 by J. McPhail.
NUMBER 172 13

FIGURE 7—Dentition of Cheirodon mitopterus, paratype,

USNM 208513, female, 35.5 mm; Panama, Code Province,

Rio Code del Norte basin, Rio Tucue at junction of river

and road between Tucue and Tambo, collected 28 August

1972 by J. McPhail.

median cusp; all teeth in a single series (Figure 7).

Maxillary with 2-3, usually 2, teeth; premaxillary

with 5 teeth in all specimens. Dentary with 8 to

10 teeth, broader than those of maxillaries, becoming smaller toward rictus, row ending in a

small conical tooth. No teeth on vomer, palatines,

or pterygoids.

Fontanel moderate, that part anterior to epiphyseal bar about as long as greatest width of

fontanel posterior to bar. Gill rakers moderate,

18-20, usually 19. Circumorbital bones well ossified, infraorbital 3 wide, contacting the preopercle

ventrally, and with a narrow naked posterior area.

Scales moderately large, cycloid, with concentric circuli, and about 4-6 radii on exposed posterior field. Lateral line complete, with up to 37-

38 perforated scales and a short fleshy tubular

extension onto middle caudal-fin rays. Lateral line

with a ventral curve on side of body, approaching

midline at end of caudal peduncle. Lateral scales

34-36, usually 35 (often irregular and difficult to

count with accuracy); scales above lateral line 5-6,

usually 5; scales below lateral line 3-4, usually 4.

Predorsal scales about 11-12. Scale sheath at base

of anal fin of about 6 scales. Axillary scale present

dorsal to pelvic-fin insertion. Caudal fin with scales

arranged much as in C. dialepturus, that is, with

approximately 2 rows of irregular scales caudad to

the more regular scales of the caudal peduncle,

extending slightly over caudal fin (some of those

near midline attached to fin membrane). Usually

with each of 2 or 3 posteriormost scales at midline

longer than deep.

Dorsal fin with ii,9 in all specimens. Dorsal-fin

origin anterior to anal-fin origin, posterior to

pelvic-fin origin, nearer eye than caudal-fin base.

Distance from tip of snout to dorsal-fin origin 40.6-

68.4. In adult males second or third rays extend

as a short filament, in females second or third ray

of dorsal fin longest; in both sexes posterior rays

shorter, forming a slightly rounded posterior margin to fin; length of longest ray 23.2-39.8, holotype

damaged.

Anal fin with 4 unbranched rays and 17-19,

usually 18, branched rays. First unbranched ray

not always visible except in radiographs. Origin of

anal fin posterior to midpoint of standard length

49.7-85.0. Fourth through eighth or ninth anal

rays longer, with successive posterior rays shorter,

forming an abruptly protruding fin margin anteriorly and a straight or slightly convex margin

jjosteriorly. Dorsally recurved anal-fin hooks present in adult males only; these occur on posterior

unbranched ray and on posterior branches of first

through seventeenth (1 specimen, holotype), first

through fifteenth (1 specimen not fully matured),

first through fourteenth (1 specimen), and first

through sixth (1 immature specimen) branched

rays; only 1 hook or bilateral pair of hooks per

bony ray segment; fin often fleshy around hooks.

Posteriormost branched anal-fin rays straight, not

bending anteriad.

Pectoral fin with 1 unbranched ray and 11-12,

usually 11, branched rays. Pectoral fins of males

reach slightly beyond pelvic-fin origin; in females

pectoral fins reach just anterior to or to pelvic-fin

origin. Distance from tip of snout to dorsal end

of pectoral-fin base 18.7-30.2, and length of pectoral

fin from base to tip of longest ray 15.0-32.5.

Pelvic fin with 1 unbranched ray and 7-8, usually

7, unbranched rays, distal tip in males reaching

to fifth branched anal ray, in females distal tip

reaching from anterior to or just to anal-fin origin.

Distance from tip of snout to pelvic-fin origin 34.9-

60.0; pelvic-fin length 15.3-30.1. In males, un-
14

branched pelvic-fin ray extended as a filament. In

males, unbranched ray without hooks, first through

seventh (2 specimens, including holotype), first

through sixth (1 specimen, immature), and second

through sixth (1 specimen, immature) branched

rays with antrorse hooks, usually on the ventral

surface o£ the rays; only 1 hook present per bony

ray segment.

Caudal fin with 10/9 principal caudal rays in

all specimens; fin forked, not split to base. Caudal

fin symmetrical and without hooks. Ventral procurrent rays of males often somewhat longer than

dorsal procurrent rays.

Precaudal vertebrae 16-17, usually 17. Total vertebrae 34-36, usually 35.

Color in alcohol: Ground color light cream

brown. No humeral spot; area at "pseudotympanum" darker than sides of body. Small melanophores present on head, tip of snout, and nape;

also present on scale margins of back, more numerous dorsally, forming reticulate pattern above

midline. Midline with groups of melanophores

forming a diffuse lateral stripe internal to scales.

In some specimens melanophores less numerous

or absent along sides above and below melanophores of midline. Often a few melanophores,

more numerous anteriorly, in a diffusely defined

lateral stripe or stripes just below midline; in more

heavily pigmented specimens there is a loose reticulate pattern instead of the diffuse lateral stripes.

A few scattered melanophores present between analfin base and midline, sometimes following myomere

junctions. Silvery lateral stripe present in recently

preserved specimens. Caudal spot present as an

elongate oval area of dense melanophores extending onto middle caudal-fin rays. On pectoral fins

melanophores present along first through fifth rays,

absent on others. A few melanophores present along

first through fifth pelvic-fin rays. Anal-fin rays with

small melanophores along rays (occasionally on

interradial membrane), more numerous proximally

and distally. Dorsal fin with small melanophores

along fin rays. Caudal fin with melanophores along

rays and on interradial membranes, concentrated

on dorsal rays of dorsal lobe and ventral rays of

ventral lobe, virtually absent from middle caudalfin rays (excepting caudal spot). In some specimens

preserved for a period of years, melanophores on

fins no longer visible.

Color in life: The following description has

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

been supplied to us by Dr. J. D. McPhail: Back

light green shading to bright silver on sides and

belly. Reticulate scale pattern of back not conspicuous. Between anal-fin base and midline, sides

translucent with a silvery coat. Two bright silver

areas (almost gold in some lights) on caudal fin

above and below oval caudal spot. Fins, except for

pectoral, with white tips, otherwise transparent.

Anal fin with microscopic flecks of red in males.

Range: Cheirodon mitopterus is found only in

the Rio Code del None basin, an Atlantic drainage of central Panama.

According to McPhail (personal communication), the Rio Code del Norte has a fauna somewhat differentiated from the rest of Panama. The

taxonomy of the forms in this river needs to be

examined in terms of broad-based studies of related forms throughout Central and South America.

Whether or not other forms in the river system

will prove to be as distinct as C. mitopterus remains to be seen.

COMPARISONS.—Cheirodon mitopterus is allopatric in regard to all known Central American

cheirodontins. Based on tooth morphology, it

seems closest to C. dialepturus and C. gorgonae but

differs as described under those species. Cheirodon

mitopterus differs from C. affinis and C. terrabae

in caudal-peduncle squamation, in having symmetrical premaxillary teeth, an enlarged median cusp

on each dentary tooth, fewer scales below the lateral line (usually 4 compared to 5 or 6 in those

species), fewer branched anal-fin rays (usually 18

compared with as many as 21-23 in those species),

and more vertebrae (usually 35-36 vs. 31-34).

Cheirodon mitopterus differs from Carlana eigenmanni in numerous features including dentition,

number of branched anal rays, gill rakers, and in

caudal squamation. Cheirodon mitopterus differs

from Phenagoniates macrolepis in having more

than 3 cusps on the teeth and a shorter anal fin,

in caudal squamation, and in numerous other characters listed under C. macrolepis.

ETYMOLOGY.—From the Greek mitos (thread)

and pteros (fin), in reference to the threadlike extensions of the dorsal and pelvic fins.

Cheirodon gorgonae Evermann and Goldsborough

FIGURES 8, 9, 10

Cheirodon gorgonae Evermann and Goldsborough, 1909:99
NUMBER 172 15

[original description; Gorgona, Panama], fig. 1 [holotype

figured], 100, fig. 3 [teeth figured].

Cheirodon insignis (not Stcindachncr).—Eigenmann, 1915:69

[Eigenmann placed Cheirodon gorgonae Evermann and

Goldsborough in the synonymy of Cheirodon insignis Steindachner].

Compsura gorgonae.—Meek and Hildebrand, 1916:274 [description], fig. 2 (teeth figured) Eigenmann, 1922:128 [generic

allocation]—Breder, 1927:117 [listed; variation, range],

163 [in key].—Jordan, Evermann, and Clark, 1930:96

[listed].—Hildebrand, 1938:250 [discussion; range].—Evans,

1952:44 [listed from Chilibre].—Gosse, 1966:8 [listed from

Panama Province].—Miller, 1966:784 [listed].—Bussing, 1967:

211 [in part, listed], 214 [in part, compared with Pseudocheirodon] Lopez, 1972:93-129 [in part, confused with

C. dialepturus and C. mitopterus], 119, fig. 4 [caudal

squamation figured inaccurately].

MATERIAL EXAMINED

(All specimens from Panama)

•USNM 64094, holotype, a juvenile 22.1 mm, Canal Zone,

Gorgona, from small seepage pool below reservoir. A. H.

Jennings, 7 Feb. 1908.

•USNM 127086, paratypes, with same data as holotype; 3

specimens 19.4 to approximately 23.0 mm (specimen damaged). Previously Bureau of Fisheries No. 5421.

•USNM 64095, paratypes, with same data as holotype; 4

specimens 19.8-21.3 mm.

•USNM 78691, Rio Frijoles, Frijoles, S. E. Meek and S. F.

Hildebrand, 14 Mar. 1911; 17 specimens 22.8-26.6 mm.

•USNM 78687, Gatun River at Monte Lirrio, S. E. Meek and

S. F. Hildebrand, 27 Mar. 1911; 41 specimens 22.1-25.5 mm.

•USNM 208543, creek about 2 mi E of Nueva Emperador

road, H. Loftin, E. Tyson, C. Kupfer, 17 Aug. 1962; 17

specimens 24.6-28.1 mm (HL-163; A7-2).

USNM 64773, sluggish stream emptying into dammed-up lake

at Gatun, A. H. Jennings, 15 Aug. 1909; 1 specimen 21.8

mm.

USNM 109254, Madden Lake, Boqueron River (above lake in

current), S. F. Hildebrand, 12 Feb. 1935; 4 specimens 26.0-

29.0 mm.

USNM 109255, Rio Cocoli, Miraflores Lake, S. F. Hildebrand,

2 Apr. 1937; 5 specimens 24.0-26.3 mm.

USNM 208542, creek about 5 mi W of El Llano on road, H.

Loftin and C. Kupfer, 17 Mar. 1962; 4 specimens 26.7-29.4

mm (HL-132; P6-2).

USNM 78694, Reservoir Creek, Gorgona, S. E. Meek and

S. F. Hildebrand, 29 Mar. 1912; 4 specimens 28.0-28.4 mm

(1 cleared and stained).

ANSP 99865, creek into Rio Caimito 8 mi from Chorrera on

Nuevo Emperado road, H. Loftin, E. Tyson, C. Kupfer,

17 Aug. 1962; 87 specimens, 14.8-24.5 mm (HL-162; P8-4).

ANSP 104039, Rio Perequete at bridge on IAH E of Capira,

H. Loftin, 24 Mar. 1962; 9 specimens 23.3-26.0 mm (HL139; P8-6).

The type series of C. gorgonae includes only

juveniles (designated as such in the morphometric

tables). For this reason, the range for adults from

several lots is given in the description and in the

tables; adults are from USNM 78687, 5 males and 5

females; USNM 70691, 3 males and 1 female; and

USNM 208543, 6 males and 4 females. In the description, the range for a given morphometric

character is given first for juveniles, then for the

adults.

FIGURE 8.—Cheirodon gorgonae, USNM 208543, male, 26.0 mm; Panama, Rio Chagres basin,

creek 2 miles east of Nuevo Emperador Road, collected 17 August 1962 by H. Loftin, E. Tyson,

and C. Kupfer.
16

Also, in stating number of "cotypes," Evermann

and Goldsborough (1909) made an error in indicating number of specimens in the USNM collections (now including the Bureau of Fisheries

specimens). On subtracting 4 specimens sent to

other institutions from the original 11, they indicate a remainder of 8 specimens—this should be 7.

DESCRIPTION.—Standard length of examined

specimens 19.4-21.3; 22.9-28.4 mm. Body elongate,

compressed laterally; greatest body depth 25.7-

36.7; 33.4-37.8. Predorsal body profile somewhat

convex with a slight concavity at nape; concavity

deepest at posterior termination of supraoccipital

spine. Body profile straight or slightly convex between posterior dorsal-fin base termination and anterior base of adipose fin. Between posterior

adipose-fin base and dorsal procurrent caudal-fin

rays, body profile slightly convex or straight. Distance from eye to dorsal-fin origin 36.2-39.8; 38.0-

43.2; distance from dorsal-fin origin to end of caudal peduncle 49.3-54.6; 50.2-54.5. Ventral body

profile gently rounded from jaws to anus; steepest

inclination ventral to jaws. Ventral body profile

protrudes its greatest distance just anterior to

pelvic-fin origin. Body profile along anal-fin base

straight or slightly convex. Between posterior termination of anal-fin base and ventral procurrent

caudal-fin rays, body profile straight or slightly

concave. Caudal peduncle depth 11.1-14.2; 12.0-

14.3; peduncle length 14.0-15.1; 13.6-17.1.

Head length 25.3-27.3; 23.5-26.3. Eye diameter

10.4-11.9; 9.2-10.8. Snout length 5.4-5.8; 5.3-5.8.

Least bony interorbital 7.0-8.3; 6.5-8.1. Maxillary

relatively long, sloping ventrally and posteriorly,

forming an angle of about 50-60 degrees to longitudinal body axis; upper-jaw length 7.6-8.3; 7.4-

8.0. All jaw teeth in a single series; teeth usually

with middle cusps enlarged, forming a rounded

cutting edge. Maxillary teeth 1-3, usually 2, usually

with about 8 cusps, teeth broader than premaxillary teeth. Premaxillary teeth 5 in all specimens

(holotype damaged), usually with 9 cusps. Dentary

with about 9 teeth, anterior 5 teeth larger (with

7-8, usually 8, cusps) and posterior teeth tapering

in size (the smallest has 3-5 cusps). No teeth present on vomer, palatines, or pterygoids.

Fontanel moderate, that part anterior to epiphyseal bar about as wide posteriorly and about

as long as widtli of fontanel immediately posterior

to bar. Gill rakers moderate, 16-20, usually 18-19

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(holotype and juveniles not counted). Circumorbital bones well ossified, infraorbital 3 wide, contacting the preopercle ventrally and with a narrow

naked area posteriorly. Infraorbital 2 not contacting the preopercle as indicated by Lopez (1972).

Scales moderately large, cycloid with concentric

circuli and about 2-5 radii on the exposed posterior

field. Perforated lateral-line scales 8-18, usually

11-13 (holotype with 10), lateral line with a slight

ventral curve. Lateral scales 32-33 (holotype with

33); scales above lateral line 5 (in some other lots

examined, there were 6 scales above lateral line);

scales below lateral line 4 in all specimens. Scale

sheath at base of anal fin of about 5-7 scales.

Axillary scale present dorsal to pelvic-fin origin;

caudal fin without scales except as described below.

Dorsal fin with ii,9 in all specimens. Dorsal-fin

origin anterior to anal-fin origin, posterior to

pelvic-fin insertion, nearer eye than caudal-fin base.

Distance between tip of snout and dorsal-fin origin

49.5-53.9; 51.4-56.8. Second or third ray of dorsal

fin longest with posterior rays shorter, forming a

slightly rounded posterior margin to fin; length of

longest ray, 29.8; 27.6-32.4 (holotype and all but

1 paratype damaged).

Anal fin with 4 unbranched rays and 15-19,

usually 16-17, branched rays. First unbranched ray

FIGURE 9.—Dentition of Cheirodon gorgonae, USNM 78694,

male, 28.3 mm; Panama, Canal Zone, Reservoir Creek,

Gorgona, collected 29 March 1912 by S. Meek and S. Hildebrand.
NUMBER 172

not always visible except in radiographs. Anal-fin

origin posterior to midpoint of standard length

63.1-65.6; 62.2-67.2. Fourth through eighth anal

rays longer than other anal-fin rays; each posterior

successive ray somewhat shorter than its predecessor, forming a protruding fin margin anteriorly

and a concave margin posteriorly. Adult males

with dorsally recurved hooks present on posteriormost branched ray and posterior branches of first

through eleventh (1 specimen), thirteenth (2 specimens), fourteenth (7 specimens), or fifteenth (4

specimens) branched rays. Hooks, single or, usually,

in bilateral pairs; only 1 hook or pair of hooks

per bony ray segment; fin often fleshy about hooks.

17

Posterior 8 or 9 branched anal-fin rays straight or

slightly curved caudad, not curved anteriorly.

Pectoral fin slightly pointed or rounded, with 1

unbranched ray and 9-11, usually 10, branched

rays (holotype damaged). Pectoral fins extend from

just anterior to slightly beyond pelvic-fin origin.

Distance from tip of snout to dorsal end of pectoralfin base 26.7-28.6; 24.2-28.8 and length of pectoral

fin from base to tip of longest ray 21.8; 20.0-25.4

(holotype and all but one paratype damaged).

Pelvic fin with 1 unbranched ray and 7-8, usually

7, branched rays, distal tip reaching from just

anterior to or to anal-fin origin. Distance from tip

of snout to pelvic-fin origin 46.9-49.0; 45.0-49.8;

FIGURE 10 Caudal fin scalation of Cheirodon gorgonae, USNM 208543, female, 28.0 mm; Panama, creek about 2 miles east of Nueva Emperador Road, collected 17 August 1962 by H. Loftin,

E. Tyson, and C. Kupfer.
18

pelvic-fin length 16.0-18.0; 16.5-21.4 (holotype

damaged). In males first unbranched pelvic ray

usually without hooks (hooks were present on 1

specimen), first through sixth (2 specimens) or

seventh (12 specimens) branched rays with antrorse

hooks on ventral surface of ray; only 1 hook present

per bony ray segment.

Caudal fin with 10/9 principal caudal rays in

all specimens (holotype damaged); fin forked, not

split to base. Upper and lower lobes usually equal

in size, occasionally a large male with slightly

larger lower caudal lobe. Dorsal and ventral procurrent caudal-fin rays usually protrude abruptly

from caudal peduncle and each ventral ray is equal

in size to its dorsal counterpart. No hooks on caudal

fin. Lower caudal lobe of both sexes with a series

of modified scales forming 3 pouches (Figure 10).

Holotype damaged and missing these scales but

scale pockets indicate presence of pouch scales;

all paratypes possess pouch scales.

Precaudal vertebrae 15-16, total vertebrae 33-34,

usually 33.

Color in alcohol: Ground color pale brown or

tan. Melanophores form a reticulate pattern on

scale pockets along back and extend over nape

and top of head; these melanophores more numerous in some populations, less in others. MelanoFIGURE 11.—Caudal fin scalation of Cheirodon heturura, holotype, FMNH 57825, male 28.4 mm; Brazil, Quimadas, Rio

Itapicuru, collected 2 March 1908 by J. Haseman.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

phores less numerous below midline, a few scattered

above anal-fin base, often along myomere junctions; belly usually unpigmented. Humeral spot

absent, but some darkening of tissues in area of

"pseudotympanum." Melanophores concentrated at

posterior termination of caudal peduncle, forming

a caudal spot. Dorsal, caudal, anal and, usually,

pelvic fins with few scattered melanophores along

fin rays, pectoral fins without pigment; otherwise

all fins hyaline.

Range: Existing collections of C. gorgonae indicate a limited distribution near the center of

Panama, where it occurs in both the Pacific and

Atlantic drainages, in Colon and western Panama

Provinces.

COMPARISONS.—Cheirodon gorgonae can be distinguished from all other Central American cheirodontins by a combination of the following three

characters: modified caudal-fin scales and peduncle

scales, forming pouches; symmetrical dentary teeth

with enlarged median cusps and absence of antrorse

hooks on the caudal fin. In the past C. gorgonae

has been confused with C. dialepturus (see the

discussion above of the latter). Cheirodon gorgonae

does seem close to C. dialepturus but differs in

having caudal scale pouches, lacking caudal-fin

hooks, and in having fewer maxillary teeth (usually

2, compared with a mode of 3 in C. dialepturus).

Cheirodon gorgonae differs from C. mitopterus

in many ways, including caudal peduncle squamation, color pattern, and meristic characters, including number of pectoral-fin rays (9-11, usually 10,

vs. 11-12, usually 11, in C. mitopterus) and total

number of vertebrae (32-34, usually 33, vs. 34-36,

usually 35, in C. mitopterus). Cheirodon gorgonae

differs from C. affinis and C. terrabae in having

symmetrical premaxillary teeth, enlarged median

cusps on the dentary teeth, forming rounded cutting edges, and a relatively longer, more slender

maxillary. Cheirodon gorgonae differs from Phenagoniates macrolepis and Carlana eigenmanni in

dentition, caudal scale modifications, and numerous

meristic characters as described.

Cheirodon gorgonae exhibits sexual dimorphism

by having retrorse hooks on the anal and pelvic

fins in males. We could find no geographical or

seasonal variation in this character. There is some

slight variation in the formation of the caudal-fin

pouches regarding size and placement of individual

scales.
NUMBER 172

Cheirodon affinis (Meek and Hildebrand)

FIGURES 12, 13, 14

Cheirodon insignis (not Steindachner).—Evermann and Goldsborough, 1909:98 [brief description], 100, fig. 2 [teeth figured].—Eigenmann, 1915:69 [in part].

Pseudocheirodon affinis Meek and Hildebrand, 1916:pl. XVIII

[photograph], 275 [new genus and species; original description; Rio Gatun, Panama] Eigenmann, 1922:129 [listed].—

Breder, 1925:143 [listed from Rio Tapia, Panama] Breder,

1927:118 [listed; variation; range], 163 [in key] Jordan, Evermann, and Clark, 1930:96 [listed] Grey, 1947:

179 [listing of types (see below)].—Hildebrand, 1938:251

[brief description; range] Gosse, 1966:8 [listed from tributary of Rio Bayano] Miller, 1966:784 [listed; range].—

Bussing, 1967:214 [compared with C. terrabae and with "C

gorgonae," a mixture of C. gorgonae and C. dialepturus],

236 [listed], 241, fig. 1 [photograph].—Gery, 1972:73 [brief

discussion in comparison with undescribed characid from

Ecuador].—L6pez, 1972:93-129 [compared with "C. gorgonae"

(a mixture of C. dialepturus, C. 7nitopterus, and C. gorgonae) and C. terrabae; variation of several characters

(discussed below); distribution], 119, fig. 3B [teeth figured].

MATERIAL EXAMINED

(All specimens from Panama)

•FMNH 8944, holotype, a male 29.5 mm: Rio Gatun, Monte

Lirio, S. E. Meek and S. F. Hildebrand, 28 Mar. 1911.

•USNM 78668, Gatun River, Monte Lirio, S. E. Meek and

S. F. Hildebrand, 28 Mar. 1911; 39 specimens 25.1-37.8 mm.

USNM 208519, Code Province, Rio Grande basin, Rio

Churube at bridge on I AH 13 mi E of Nata, H. Loftin

and C. Kupfer, 25 Feb. 1962; 34 specimens 22.3-32.1 mm

(5 cleared and stained) (HL-118; P10-3).

USNM 208520, Panama Province, Rio Bayano Basin, creek

about 5 mi W of El Llano on road, H. Loftin and C.

Kupfer, 17 Mar. 1962; 36 specimens 27.2-39.2 mm (2 cleared

and stained) (HL-132; P6-2).

USNM 62943, small ditch of "pure" running water at

Tabernilla, Canal Zone, A. H. Jennings, 24 July 1908; 6

specimens 24.3 29.7 mm.

19

USNM 78667, Gatun River at Mitchelville, S. E. Meek and

S. F. Hildebrand, 27 Mar. 1911; 36 specimens 28.3-37.9 mm.

USNM 208545, Canal Zone, small ditch of "pure" running

water at Tabernilla, A. H. Jennings, 24 July 1908; 4 specimens 24.7-25.8 mm.

USNM 78657, Limon Creek, Alhajuela, S. E. Meek and S. F.

Hildebrand, 26 Feb. 1911; 9 specimens 24.4-35.2 mm.

USNM 78664, Gorgona Reservoir Creek, Gorgona, Canal

Zone, S. E. Meek and S. F. Hildebrand, 29 Mar. 1912; 8

specimens 34.8—42.8 mm.

USNM 78681, Rio Missimbi, Empire, Canal Zone, S. E. Meek

and S. F. Hildebrand, 8 Feb. 1911; 12 specimens 28.4-34.9

mm.

USNM 78684, Trinidad River, Agua Clara, Canal Zone, S. E.

Meek and S. F. Hildebrand, 10 Mar. 1911; 24 specimens

24.5-35.3 mm.

USNM 208544, Panama Province, Rio Bayano basin, creek

about 2<4 mi W of El Llano road, H. Loftin and C. Kupfer,

17 Mar. 1962; 10 specimens 29.3-34.0 mm (HL-131; P6-3).

USNM 208518, Veraguas Province, Rio San Pedro basin, creek

at bridge 12 mi W of Santiago on road to Sona, H. Loftin,

E. Tyson, R. Yerger, 28 Jan. 1962; 49 specimens 18.6-29.3

mm (2 cleared and stained) (HL-105; P16-9).

USNM 208548, Code Province, Rio Grande Basin, Rio Code

at road just outside La Pintada, H. G. Loftin, 23 Mar.

1962; 59 specimens 22.2-35.0 mm (HL-135; P10-2).

USNM 208550, Rio Santa Maria Basin, Rio Estero Salado

(coastal stream) on IAH about 5 mi W of Aguadulce, H.

Loftin and E. Tyson, 15 Oct. 1961; 11 specimens 24.3-29.4

mm (HL-24; PI 1-2).

USNM 208549, Veraguas Province, Rio San Pedro basin,

river at bridge about 17 mi W of Santiago on road to

Sona, H. Loftin E. Tyson, R. Yerger, 28 Jan. 1962; 12

specimens 21.8-27.7 mm (HL-104; P16-10).

USNM 208551, Code Province, Rio Grande basin, Rio Anton

at town of Anton on IAH, H. Loftin and E. Hcslop, 11

Mar. 1962; 2 specimens 29.9-31.2 mm (HL-I20; P9-21).

USNM 208515, Code Province, Rio Grande basin, swampy

creek at IAH about 2 mi E of Nata. H. Loftin and E.

Tyson, 15 Oct. 1961; 6 specimens 29.7-35.7 mm (2 cleared

and stained) (HL-21; PI0-5).

MCZ 45827, "Jesus Christ Stream" trib. of Chagres River

near Gamboa, 9°7'N, 79°42'W. I. Rubinoff, et al., 6 Apr.

1967; 1 specimen 30.0 mm.

*

& - ^

FIGURE 12 Cheirodon affinis, holotype, FMNH 8944, male, 29.5 mm; Panama Rio Gatun, Monte

Liria, collected 28 March 1911 by S. Meek and S. Hildebrand.
20

USNM 210991, two collections mixed together, Code Province,

stream at El Roble at IAH or Veraguas Province, Rio San

Pedro at San Pedro on IAH, M. and W. Bussing, 27 January 1971; 3 specimens 26.5-32.2 mm.

Meek and Hildebrand (1916) listed the holotype

number as FMNH 8941 and stated that it was

"38 mm in length." Grey (1957) stated that the

above number is an error and that the actual

number is 8944. Mr. Loren P. Woods has kindly

reexamined the catalog and confirmed that FMNH

8944, the number in the bottle, is indeed correct.

The holotype is 29.5 mm in SL.

Grey (1947) listed 261 paratypes from several

areas in Panama. We do not accept type status

for any of these fishes as Meek and Hildebrand

(1916), although using more than 1 specimen for

their description, listed only the holotype, adding

that "numerous specimens were preserved."

Tables include specimens used in the following

description and 10 specimens each USNM 208519

and USNM 208520.

DESCRIPTION.—Standard length of examined

specimens 22.5-38.8 mm. Body elongate, compressed laterally; greatest body depth 38.0—42.1.

Predorsal body profile convex with a slight concavity at nape; concavity deepest at posterior termination ot supraoccipital spine. Body profile slightly

convex between posterior dorsal-fin base and adipose fin; posterior to adipose fin body profile with

an upward slope to procurrent caudal-fin rays.

Distance from eye to dorsal-fin origin 38.9-41.9;

distance from dorsal fin to end of caudal peduncle

53.6-56.0. Ventral body profile rounded to anus;

steepest inclination ventral to jaws. Ventral body

profile protrudes ventrally its greatest distance just

anterior to pelvic-fin insertion. Body profile straight

or slightly curved along anal-fin base; at posterior

anal-fin termination profile concave to procurrent

caudal-fin rays. Caudal-peduncle depth 12.3-13.7;

peduncle length 11.9-14.6.

Head length 23.4-25.4. Eye diameter 9.3-10.6.

Snout 5.3-6.2. Least bony interorbital width 8.2-

9.0. Maxillary short, sloping ventrally and posteriorly, forming an angle of 45-50 degrees to

longitudinal body axis; upper-jaw length 6.2-7.1.

All teeth with 7-8, usually 8, cusps (except smallest maxillary tooth, which has 5-6 cusps, and the

smallest dentary teeth, which are quinquecuspid

or conical); all in a single series. Maxillary teeth

3, with a rounded cutting edge and with outer

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cusps equidistant from tooth base. Premaxillary

teeth 5 in all specimens with a rounded cutting

edge, cusps on anterior cutting edge begin nearer

to tooth base than cusps on posterior cutting edge.

Dentary with 7-9, usually 7-8, teeth, anterior teeth

larger, becoming smaller (usually conical) posteriorly; teeth with an even cutting edge, median

cusp not elongate, long axis of anterior dentary

teeth extending dorsoanteriorly. No teeth present

on vomer, palatines, or pterygoids.

Fontanel moderate, length of that part anterior

to epiphyseal bar slightly less than width of fontanel posterior to bar. Gill rakers moderate, 17-

20, usually 18. Circumorbital bones well ossified,

infraorbital 3 wide, contacting preopercle ventrally,

and with a naked area posteriorly.

Scales moderately large, cycloid with concentric

circuli and about 4-8 radii on exposed posterior

field. Lateral lines of holotype and specimens from

USNM 78668 incomplete, with a slight ventral

curve with 7-11, usually 9 (10 in holotype), perforated scales. In some other lots examined lateral

line variable (e.g., USNM 208520), with some specimens having 10-11 perforated scales and others in

the same lot with as many as 33 perforated scales

(a complete lateral line). This variation not present

in all populations but occurs throughout range

of this species (see discussion below). Lateral scales

FICURE 13.—Dentition of Cheirodon affinis, USNM 208519,

female, 25.6 mm; Panama, Code Province, Rio Grande basin,

Rio Churube at bridge on IAH 13 miles east of Nata, collected 25 February 1962 by H. Loftin and C. Kupfer.
NUMBER 172

32-33 (33 in holotype); scales above lateral line

5-6, usually 6; scales below lateral line 4-5, usually

5. Predorsal scales 9-12, usually 10-11. Scale sheath

at base of anal fin of about 5-6 scales. Axillary

scale present dorsal to pelvic-fin insertion; caudal

fin without scales.

Dorsal fin ii,9 in all specimens. Dorsal-fin origin

anterior to anal-fin origin, posterior to pelvic-fin

insertion, nearer eye than caudal-fin base. Distance

from tip of snout to dorsal-fin origin 51.7-54.5.

Third or fourth ray of dorsal fin longest, with

posterior rays shorter, forming a slightly rounded

posterior margin to fin; length of longest ray

30.0-30.9.

Anal fin with 4 unbranched rays and 19-21,

usually 20, branched rays. First unbranched ray

not always visible except in radiographs. Anal-fin

origin posterior to midpoint of standard length

63.6-67.5. Fourth through ninth or tenth anal-fin

rays longer, with posterior rays shorter, forming a

protruding fin margin anteriorly and a straight or

slightly concave fin margin posteriorly. Dorsally

recurved anal-fin hooks present in males only (see

discussion below) on last unbranched ray and first

21

through sixth or seventh (rarely to eleventh)

branched rays (on first through sixth in holotype);

usually only 1 hook or bilateral pair of hooks per

bony ray segment (rarely with 2 hooks on same

side of segment); fin often fleshy around hooks.

Pectoral fin with 1 unbranched ray and 10-11,

usually 11, branched rays. Pectoral fins reach anterior to or just to pelvic-fin insertion. Distance

from tip of snout to dorsal end of pectoral-fin base

24.5-26.4 and length of pectoral-fin base to tip

of longest ray 20.1-21.8.

Pelvic fin with i,7 rays in all specimens, distal

tip reaching well anterior to or just to anal-fin

origin. Distance from tip of snout to pelvic-fin

insertion 47.4-49.8; pelvic-fin length 16.9-19.1. In

males, unbranched ray and first or second through

sixth or seventh branched rays with antrorse hooks,

usually on ventral surface of ray segment (unbranched ray often without hooks); 1 hook, rarely

2, present per bony ray segment (in holotype all

pelvic rays with hooks). See below for further

discussion.

Caudal fin with 10/9 principal rays; fin not

deeply forked, not split to base; caudal-fin lobes

FIGURE 14 Cheirodon affinis, USNM 210991, female, 26.5 mm; Panama (from two collections

mixed together), Code Province, stream at El Roblc at IAH or Veraguas Province; Rio San

Pedro at San Pedro on IAH, collected 27 January 1971 by Myrna and W. Bussing; specimen

photographed in life.
22

symmetrical. Dorsal and ventral procurrent caudal

rays of equal size; ventral procurrent rays not enlarged and extended. No hooks on caudal rays.

Precaudal vertebrae 15-16, usually 16. Total

vertebrae 32-33, usually 33.

Color in alcohol: Ground color pale brown,

back and nape darker brown. Small melanophores

present on sides bordering scale pockets, more numerous dorsally, forming a reticulate pattern above

midline and sparingly across belly (not as evident

in type as in some other specimens) and loosely

following myomere junctions below midline above

anal fin. Melanophores sometimes slightly concentrated at "pseudotympanum," forming a weak

humeral spot. Melanophores usually concentrated

just dorsal to midline, forming a lateral stripe

which terminates in a large spot at end of caudal

peduncle, extending to middle caudal rays. Lateral

stripe not always pronounced but caudal spot always present. Type with silvery cheeks and with a

broad, silvery lateral stripe; this silvery coloration

more or less evident in other specimens examined.

Dorsal fin with melanophores on interradial membranes, more concentrated distally. Caudal fin with

melanophores on interradial membranes, often concentrated near middle caudal rays. Anal fin with

melanophores on interradial membranes (occasionally on rays) proximally, but melanophores lacking

on distal one-half to one-fourth of fin. Pectoral fin

with melanophores ventrally on rays, lacking on

interradial membranes. Pelvic fins with melanophores primarily on interradial membranes, concentrated distally.

Color in life: Based on 1 male and 3 females

from USNM 210991 observed alive in our aquaria

(with dark substrate), the color is as follows. Back

light yellowish-green shading to bright silver on

sides and belly. Burnished silver lateral stripe.

Reticulate scale pattern of back conspicuous. Between anal-fin base and midline, side translucent

yellowish green, with loose pattern of melanophores along myomere lines. Light yellowish areas

on caudal fin above and below broad caudal spot.

Dorsal, anal, and pelvic fins with small white tips;

all fins but pectorals yellowish, pectoral fin translucent.

Range: Collections of C. affinis are most numerous from the Canal Zone, probably reflecting

the extensive collecting in that area early in this

century. Loftin (1965) extended the known range

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of this species from east-central Panama Province

to central Veraguas Province. The species is now

known in Pacific drainages throughout central

Panama and in Atlantic drainages at least throughout the Canal Zone.

COMPARISONS.—Cheirodon afjinis is a common

species in central Panama. It is sympatric with

C. dialepturus and C. gorgonae and there has been

some confusion among these three species. Cheirodon afjinis is distinguished from the other two by

the combination of its lack of both caudal-fin hooks

and modified caudal-peduncle scales, together with

its possession of asymmetrical premaxillary teeth

and dentary teeth without an enlarged median

cusp. It differs from C. mitopterus in dentitional

characters also, as well as caudal peduncle scalation, and other characters as described in the discussion of C. mitopterus.

Cheirodon affinis is closest to C. terrabae but

apparently differs from that form in having fewer

dentary and maxillary teeth, a smaller modal number of branched anal-fin rays, and gill rakers. Most

populations of C. affinis have an incomplete lateral

line, whereas terrabae has the lateral line complete.

VARIATION.—Previously, C. affinis has been described as lacking anal-fin hooks (Bussing, 1967).

Our examination has shown that retrorse hooks

are present on the anal and pelvic fins of males

collected between the months of March and July

and are absent in those specimens collected in the

months of October and January. We were unable

to obtain collections from every month, so our

information on this variation is incomplete. Also

we had no collections from the same locality from

different seasons, and the presence or absence of

hooks could be explained as population variation.

However, numerous collections from different seasons are from adjacent localities in the Canal Zone

and these showed seasonal variation. Other than

the hooks on the fins of males, we could find no

obvious sexual dimorphism.

Bussing (1967) and Lopez (1972) discussed the

length of the lateral line in C. affinis. We have

essentially confirmed their findings, as we have

found specimens from several localities with complete or nearly complete lateral lines. We could

find no geographical or ecological correlation with

the variation in this character.

Lopez (1972) found a correlation in length of

the maxillary and geographical distribution. We
NUMBER 172 23

found no significant correlation with total upperjaw length and distribution. See our comments

on this character in the discussion of C. dialepturus.

Cheirodon terrabae (Bussing)

FIGURES 15, 16

Pseudocheirodon species.—Miller, 1966:784 [listed].

Pseudocheirodon terrabae Bussing, 1967:212 [original description; Rio Grande de Terraba, Costa Rica], 241, fig. 1

[photograph] L6pez, 1972:101 [listed from Terraba and

Pirris river basins; in her reprints Lopez has indicated that

the Turrubares basin is an error and should be Pirris

(PI)], 105 [compared to C. affinis], 117, fig. 2 [distribution

map], 121, fig. 6 [photograph], 123, fig. 7 [distribution map],

129, fig. 14 [scales].

MATERIAL EXAMINED

(All specimens from Costa Rica)

•LACM 2951, paratypes, Costa Rica: San Jose Province, tributary to Rio General, W. A. Bussing, 28 Dec. 1961; 12

specimens 30.0-47.1 mm (1 cleared and stained).

•USNM 194236, Rio Union, IAH near San Isidro, Albert A.

Greenberg, 20 Jan. 1963; 4 specimens 30.4-36.0 mm (1

cleared and stained).

•ANSP 121988, with same data as USNM 194236, 1 specimen

35.4 mm.

•CAS 16061, with same data as USNM 194236, 1 specimen

31.1 mm.

•BMNH 1973^.16;2, with same data as USNM 194236, 1

specimen 35.4 mm.

•USNM 194234, Rio Alia Nuo, near IAH, Albert A. Greenberg, 21 Jan. 1963; 2 specimens 30.2-30.4 mm.

•USNM 194220, with same data as USNM 194234; 1 specimen too damaged to measure.

•UMMZ 194214, overflow pond of Rio Grande de Terraba,

Palmar Norte, R. M. Bailey and party, June 1973; 32

specimens 39.7-50.5.

Specimens in description are included in tables.

DESCRIPTION.—Standard length of specimens examined 29.8-49.8 mm. Body deep, compressed laterally; greatest body depth 32.9-42.6. Predorsal

body profile somewhat convex with a slight concavity at nape. Between posterior dorsal-fin termination and anterior adipose-fin base, body profile

very slightly convex. Some specimens with a slight

rise in body profile at base of adipose fin; between

posterior adipose-fin termination and dorsal procurrent caudal-fin rays, body profile slightly concave. Distance from eye to dorsal-fin origin

37.2-43.0; distance from dorsal-fin origin to posterior end of caudal peduncle 51.0-58.0. Ventral

body profile rounded from jaws to anus; steepest

inclination ventral to jaws. Ventral profile protrudes ventrally its greatest distance at point just

anterior to or just posterior to pelvic-fin insertion.

Body profile along anal-fin base straight or slightly

convex; body profile concave between posterior

anal-fin termination and procurrent caudal-fin rays.

Caudal peduncle depth 11.0-13.2; peduncle length

11.3-15.0.

FIGURE 15.—Cheirodon terrabae, paratype, LACM 2951, female, 39.3 mm; Costa Rica, San Jose

Province, tributary to Rio General, collected 28 December 1961 by VV. Bussing.
24

Head length 22.8-26.6. Eye diameter 8.7-10.4.

Snout 5.4-6:5. Least bony interorbital width 7.6-

9.2. Maxillary short, sloping ventrally and posteriorly, forming an angle of 50-60 degrees to

longitudinal body axis; upper-jaw length 6.0-7.7.

Upper-jaw teeth with 6-10, usually 8-9, cusps

(except smallest maxillary teeth, which have 5-6

cusps), all in a single series. Maxillary teeth 3-6,

usually 4-5, with a rounded cutting edge and with

outer cusps equidistant from tooth base. Premaxillary teeth 5 in all specimens with a rounded cutting

edge, cusps on anterior cutting edge begin nearer

to tooth base than cusps on posterior cutting edge.

Dentary with 8-10 teeth, with 6-10, usually 8-9,

cusps, anterior teeth larger, becoming smaller posteriorly (posterior 2 teeth may be tricuspid or

conical); teeth with an even cutting edge, median

cusp not elongate, long axis of anterior dentary

teeth extending dorsoanteriorly or may be nearly

horizontal in some specimens. No teeth present

on vomer, palatines, or pterygoids.

Fontanel moderate, length of that part anterior

to epiphyseal bar slightly less than width of fontanel posterior to bar. Gill rakers moderate, 19-22,

usually 21. Circumorbital bones well ossified, infraorbital 3 wide, contacting preopercle ventrally and

with a naked area posteriorly.

Scales moderately large, cycloid, with concentric

FIGURE 16 Dentition of Cheirodon terrabae, paratype,

LACM 2951, male 32.3 mm; Costa Rica, San Jose Province,

Tributary to Rio General, collected 28 December 1961 by

W. Bussing.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

circuli and about 5-8 radii on exposed posterior

field. Lateral line complete, extending as a ventral

curve on side of body to below adipose fin, then

continuing below midline as a straight line to

caudal-fin base, sometimes continuing slightly onto

middle caudal-fin rays. Lateral-line scales approximately 33—34. Scales above lateral line 6-7, usually

6; scales below lateral line 5-6. Predorsal scales

number about 11. Scale sheath at base of anal fin

of about 3-6 scales. Axillary scale present dorsal

to pelvic-fin insertion; caudal fin scaleless.

Dorsal fin with ii,9 rays in all specimens. Dorsalfin origin anterior to anal-fin origin, posterior to

pelvic-fin origin, nearer eye than base of caudal

fin. Distance from tip of snout to dorsal-fin origin

50.0-57.6. Third or fourth ray of dorsal fin longest,

with posterior rays shorter, forming a slightly convex posterior margin to fin; length of longest ray

27.8-34.4.

Anal fin with 4 (rarely 5) unbranched rays and

19-23, usually 22 or 23, branched rays. First unbranched ray not always visible except in radiographs. Anal-fin origin posterior to midpoint of

standard length 61.3-69.5. Fourth through seventh

or eighth anal rays longer, with successive rays

shorter, making ventral margin of anal fin concave. Dorsally recurved anal-fin hooks present in

males only (hooks were not noted in original

description) on last unbranched ray and first

through eleventh to seventeenth branched rays,

usually only 1 hook or bilateral pair of hooks per

bony ray segment (rarely with 2 hooks on same

side of segment); fin often fleshy around hooks.

Pectoral fin with 1 unbranched ray and 11-12,

usually 11, branched rays. Pectoral fins reach from

just anterior to or slightly beyond pelvic-fin insertion. Distance from snout tip to dorsal end of

pectoral-fin base 24.0-29.0 and length of pectoral

fin from base to tip of longest ray 19.4-24.1.

Pelvic-fin rays i,7 in all specimens, distal end

reaching to or just anterior to anterior basal termination of anal fin. Distance from tip of snout to

pelvic-fin origin 45.1-51.1; pelvic-fin length 16.7—

21.3. In males, first through seventh branched rays

with antrorse hooks usually on ventral surface of

ray segments; 1 hook, rarely 2, present per bony

ray segment.

Principal caudal-fin rays 10/9 (1 specimen with

9/9, 1 with 9/8); fin forked, not split to base; no

hooks present.
NUMBER 172

Precaudal vertebrae 16; total vertebrae 33-34,

usually 33.

Color in alcohol: Ground color pale brown

(gray brown in freshly preserved specimens), back

and nape darker brown. Small melanophores present on sides bordering scale pockets, more numerous dorsally, forming a reticulate pattern above

midline, loosely following myomere junctions below midline above anal fin, present sparingly

across belly. Melanophores sometimes slightly concentrated at "pseudotympanum," forming a weak

humeral spot. Melanophores concentrated at end

of caudal peduncle, forming a large caudal spot

that extends along middle caudal rays. Freshly preserved material with silvery cheeks and with a

broad silvery lateral stripe; this silvery coloration

less evident in other specimens examined. Dorsal

fin with melanophores on interradial membranes.

Caudal fin with melanophores on interradial membranes, usually concentrated near middle caudal

rays. Anal fin with melanophores on interradial

membranes (occasionally on rays) proximally, but

melanophores usually lacking on distal one-half

to one-fourth of fin. Pectoral fin with melanophores

ventrally on rays, lacking on interradial membranes. Pelvic fins with melanophores primarily on

interradial membranes, concentrated distally.

Range: Bussing (1967) states that C. terrabae

has been found in the basin of the Rio Grande de

Terrabae. Lopez (1972) lists it also from the Rio

Jicote, of the Rio Pirris basin, also in Costa Rica.

Bussing in personal communication has suggested

that the Rio Alia Nuo might be Saloma Nuevo,

8°15'N, 83°13'W.

COMPARISONS.—Cheirodon terrabae is very closely

related to C. affinis of Panama. It differs from that

species in having a greater number of dentary

teeth (8-10, usually 9, vs. 7-9, usually 7, in C.

affinis), greater modal number of maxillary teeth

(3-6, usually 5-4, vs. 3-4, usually 3, in C. affinis),

and slightly greater average number of anal-fin

rays (22.1 vs. 19.6 in C. affinis). Other apparent

differences occur in the number of scales below

the lateral line and number of gill rakers (see

tables). There may also be a greater number of

hooked anal-fin rays in C. terrabae males. In some

populations of C. terrabae (e.g., LACM 2951),

there may be a modal number of 3 maxillaty teeth,

leading to possible confusion with C. affinis.

Cheirodon terrabae is distinguished from C. dia25

lepturus, C. gorgonae, and C. mitoptems in lacking

modified caudal squamation and in its possession

of asymmetrical premaxillary teeth and dentary

teeth without an enlarged median cusp. Young

Carlana eigenrnanni, which have teeth similar to

those of C. terrabae, can be quickly identified by

the much higher number of anal-fin rays; other

differences between these species may be found

under C. eigenmanni and in the tables. Number

of anal-fin rays, position of the dorsal fin, and

dentition are among the numerous differences between C. terrabae and P. macrolepis (see P. macro

lepsis description below).

Bussing (1967) remarked:

Although it has been proposed that the present genera

Odontostilbe, Holoshesthes, and Saccoderma have retained

the complete lateral line of an Odontostilbe-like ancestor

and that the general trend in minute species is toward reduction or suppression of the lateral line . . . , such is not the

case with P. terrabae. If it is assumed that P. terrabae is a

descendant species of P. affinis, then the new species has

attained a complete lateral line and certain populations of

P. affinis are actively mutating in the same direction.

To assume that one existing species is the ancestor

of another as Bussing does is presumptuous and

probably fallacious (cf. Hennig, 1966). In all probability these two species, which seem closest to one

another, had a common ancestor not too different

from either existing species. We cannot see any

basis for predicting the characters of the ancestors

in this case, especially not by the use of such apparently labile characters as the length of a lateral

line. This is the primary reason why we have not

accepted the genus Odontostilbe (see discussion

above under the generic name Cheirodon). We do,

however, fully agree with Myers (1958) that loss

of a lateral line is a general trend in small characids. We believe that it probably happened repeatedly and independently in different groups of species

and genera, but we agree with Bussing's suggestion

that occasionally the line may have reappeared,

confusing the attempt to establish relationships by

use of this character.

Carlana Strand

Carlia Meek, 1914:108 [type-species Cheirodon eigenmanni

Meek, 1912, by original designation].

Carlana Strand, 1926:54 [type-species Cheirodon eigenmanni

Meek, 1912, replacement name for Carlia Meek, 1914, preoccupied by Carlia Gray, 1854].
26

Adults with teeth along entire ventral maxillary

margin; dentary teeth broad with 9-12 cusps; number of branched anal-fin rays 26-31 (in Carlana

eigenmanni, only known species), compared with

15-23 or 45-50 in other genera in geographical

area covered here. Weak spinules present on pelvicand anal-fin rays of adult males (see discussion

below).

Meek (1914) indicated that his new genus Carlia

was closely related to Rhoadsia Fowler, which is

currently included in a subfamily Rhoadsiinae.

Eigenmann and Myers (1929) included Carlia in

Rhoadsia but stated that the species C. eigenmanni

was very likely generically distinct from other

species of Rhoadsia. Juvenile Rhoadsia have a

single series of premaxillary teeth and gain a second

row when approaching adulthood [reaching up to

168 mm according to Eigenmann and Myers

(1929)]. The maxillary in juvenile Rhoadsia has

but 2 multicuspid teeth, while the maxillary in

adults has, in addition, numerous conical teeth

along its entire ventral margin. We have 4 adult

males (as evidenced by anal- and pelvic-fin spinules)

of Carlana eigenmanni ranging from 52.8-53.9 mm,

all of which lack an outer premaxillary tooth row

but have a fully toothed maxillary. We are confident that Carlana does not have an outer row of

premaxillary teeth. According to the present classification system, this feature alone would remove

Carlana eigenmanni from Rhoadsia and place it

in the Cheirodontinae.

We have examined several specimens of Rhoadsia as follows: R. minor (CAS[IUM] 13419, holotype; CAS[IUM] 13420, 92 specimens, paratypes;

CAS[SU] 49795, 4 specimens; CAS[SU] 49796, 1

specimen); R. altipinna (FMNH 55907, 1 specimen, paratype; USNM 83543, 1 specimen; USNM

83536, 1 specimen). Adult specimens have dentary

teeth quite different from C. eigenmanni in that

they are not nearly as broad and have 4 large cusps

with 1 or 2 smaller cusps (compared to very broad

teeth with 9-12 cusps in C. eigenmanni). Also, all

adults have a well-defined outer row of 2 or 3

premaxillary teeth in contrast to C. eigenmanni,

which has only 1 tooth row.

While these differences in dentition are, according to the currently accepted system, of generic

significance, it is obvious that a complete review

of the so-called Rhoadsiinae, sensu lato (including

Parastremma Eigenmann, Rhoadsia, and possibly

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Carlana), is needed. There are many characters in

all these species that possibly indicate relationships,

including overall similarity of body and fin shape

and ontogenetic dentition changes. Despite the

fact that Carlana, because of its single row of premaxillary teeth, must currently be defined as a

so-called cheirodontin, it is possible that further

investigation will place all the species now in

Parastremma and Carlana in Rhoadsia.

Carlana eigenmanni (Meek)

FIGURES 17, 18, 19, 20, 21

Cheirodon eigenmantii Meek, 1912:70 [original description;

La Junta, Costa Rica].

Carlia eigenmanni Meek, 1914:108 [new genus; listed from

La Junta].

Carlana eigenmanni.—Strand, 1926:54 [new name for preoccupied Carlia].

Rhoadsia eigenmanni.—Eigenmann and Myers, 1929:463 [description of specimen from La Junta], pi. 73: fig. 1 [photograph].—Grey, 1947 [correction of FMNH number of

type] Bohlke, 1958:178 [compared with Hyphessobrycon

tortuguerae; discussion of generic status].—Miller, 1966:785

[listed from Lake Nicaragua to Atlantic slope of Costa Rica

fide Rivas and Bussing].—Bussing, 1967:236 [listed from

Costa Rica].—Astorqui, 1971:20 [listed from Rio Tisla and

Rio Osayo, Nicaragua; description].

MATERIAL EXAMINED

•FMNH 7683, holotype, a male 52.8 mm, Costa Rica: La

Junta, S. E. Meek, Apr. 1912.

•FMNH 43150, Costa Rica: La Junta, S. E. Meek, 8 Apr.

1912; 1 specimen 56.4 mm.

•FMNH 7861, Costa Rica: La Junta, S. E. Meek, 17 Apr.

1913; 3 specimens 46.4-53.9 mm.

•USNM 74240, Costa Rica: La Junta, S. E. Meek; 1 specimen

53.6 mm. (maxillary, premaxillary and dentary of right

side cleared and stained).

*CAS(SU) 61380, Nicaragua: Rio Frio, about 1.5 mi above

San Carlos, in bayou with sluggish current, G. S. Myers,

12 Feb. 1963; 4 specimens 21.0-26.9 mm (1 cleared and

stained).

•CAS(SU) 61369, Nicaragua: Rio Frio, 1.5 mi above San

Carlos, 0.4 mi above Esperanza, small sluggish stream into

Rio Frio, G. S. Myers; 8 specimens 21.6-35.8 mm.

•GCRL 5126, Nicaragua: Rio Osayo at IAH, 9 km N of Costa

Rica border, L. R. Rivas and I. Astorqui, 13 July 1960; 2

specimens 40.0-43.1 mm.

•GCRL 5125, Nicaragua: Rio Isala at road from Managua

to Juigalpa, 8 km W of Juigalpa, L. R. Rivas and I.

Astorqui, 28 June 1960; 1 specimen 37.2 mm.

Meek (1912) in describing Cheirodon eigenmanni listed 1 specimen, FMNH 7583, 67 mm;
NUMBER 172 27

collected April 1912 at La Junta; subsequently

(1914), when he placed the species in Car Ha, he

listed 6 specimens from the type-locality. Grey

(1947) corrected the number on the type to 7683

and listed 1 unnumbered paratype. We presume

that the specimen Grey considered as paratype is

FMNH 43150, because it is the only other FMNH

specimen collected April 1912. However, since

Meek mentioned no paratypes in the original description we do not accept paratype status for any

specimens.

Eigenmann and Myers (1929) listed a specimen

73.0 mm from La Junta, coll. Meek, and stated

that it had minute hooks on the anal-fin rays. We

have been unable to locate their specimen after

searching the CAS, FMNH, and USNM collections.

Oft*

FIGURE 17.—Juvenile Carlana eigenmanni, CAS(SU) 61380, 26.8 mm; Nicaragua, Rio Frio, about

ly2 miles from San Carlos, Lake Nicaragua, collected 12 February 1963 by G. Myers.

FIGURE 18.—Pre-adult Carlana eigenmanni, GCRL 5125, 32.7 mm; Nicaragua, Rio Isala at road

from Managua to Juigalpa, 8 kilometers west of Juigalpa, collected 28 June 1960 by L. Rivas

and I. Astorqui.
28

In the following description, proportions for

juveniles are given first, followed by adults, separated by a semicolon. In listing meristics, counts

for juveniles and adults are combined. Juveniles

in this case are arbitrarily defined by the presence

of only 2 maxillary teeth and consist of 6 specimens from CAS(SU) 61369 and 4 specimens from

CAS(SU) 61380. All specimens are included in

tables.

DESCRIPTION.—Standard length of specimens examined 21.0-29.8; 34.3-56.4 mm. Body elongate,

compressed laterally; greatest body depth 32.0-

39.5; 35.8-41.2. Predorsal body profile convex or

slightly convex with a slight concavity at nape.

At anterior dorsal-fin base, body profile angles

ventrally to posterior termination of dorsal-fin base.

Body profile straight or somewhat convex between

posterior dorsal-fin base and adipose-fin base. Posterior to adipose fin, body profile straight or slightly

concave to dorsal procurrent caudal-fin rays. Distance from eye to dorsal-fin origin 38.9-41.5; 39.6-

41.2; distance from dorsal origin to end of caudal

peduncle 49.0-52.4; 51.9-56.3. Ventral profile in

juveniles rounded to anus, steepest inclination ventral to jaws; profile protruding ventrally its greatest distance midway between pelvic-fin insertion

and anal-fin origin. Ventral profile in adults somewhat convex to a vertical from pectoral-fin base,

then becomes more convex to pelvic-fin base; posterior to pelvic-fin base, body profile convex to

anal-fin origin. Body profile along anal-fin base

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

straight or slightly concave in juveniles, straight

or somewhat convex in adults; between posterior

anal-fin termination and ventral procurrent caudalfin rays, body profile concave. Caudal peduncle

depth 8.3-10.1; 9.5-11.6; caudal peduncle length

8.7-10.9; 9.5-10.5.

Head length 24.9-27.6; 23.1-25.2. Eye diameter

10.8-12.7; 8.5-10.8. Snout length 4.9-6.9; 6.1-7.1.

Least bony interorbital width 7.2-9.0; 8.4-8.9.

Maxillary short in juveniles, long in adults, sloping ventrally and posteriorly, forming an angle

of about 50-60 degrees to longitudinal body axis.

Upper-jaw length 7.5-8.2; 8.6-11.9. All teeth uniserial. Maxillary of juveniles with 2 proximal teeth

having 5-8 cusps (median cusps may be slightly

enlarged); maxillary of adults with 2 proximal

teeth as in juveniles but also with up to 8 more

conical teeth along its ventral margin. Premaxillary

teeth with anterior cutting edge beginning nearer

to tooth base than cusps on posterior cutting edge;

apices of teeth angled toward midline in adults.

Dentary teeth 7-9, broad, with 8-12 cusps (except

posteriormost 2 or 3 small teeth with 3 to 5 cusps);

cutting edge straight or slightly rounded with

median cusps broader, becoming narrower and

smaller laterally. No teeth on vomer, palatines,

or pterygoids.

Fontanel moderately large in juveniles, that part

anterior to the epiphyseal bar a little more than

two-thirds length of fontanel posterior to bar; in

adults, that part anterior to bar about one and

FIGURE 19.—Adult Carlana eigenmanni, holotype, FMNH 7683, male, 52.8 mm; Costa Rica, La

Junta, collected April 1912 by S. Meek.
NUMBER 172 29

one-half times as long as width of fontanel immediately posterior to bar. Gill rakers moderate, 14-

18, usually 16. Circumorbital bones well ossified,

in juveniles iniraorbital 3 with a broad naked area

ventrally and posteriorly, in adults it contacts preopercle below but with a naked area posteriorly.

Scales moderately large, cycloid with concentric

circuli and 0-5 radii on exposed posterior field.

Lateral line incomplete with 5-9, usually 8-9, perforated scales; lateral line with a slight ventral

curve. Lateral scales 35-41, usually 38 (holotype

with 38); these scales not always regularly arranged, therefore counts not always accurate. Scales

above lateral line 8-9, usually 8; scales below lateral

line 7-8, usually 7. Predorsal scales about 17. Axillary scale present dorsal to pelvic-fin origin; caudal

fin scaleless; 6 or 7 scales forming a sheath along

anterior base of anal fin.

Dorsal fin with 2 unbranched rays and 8-9,

usually 9, branched rays. Dorsal-fin origin anterior

to anal-fin origin, posterior to pelvic-fin origin,

nearer eye than caudal-fin base. Distance from tip

of snout to dorsal-fin origin 53.4-56.7; 52.0-55.0.

Third or fourth ray of dorsal fin longest, with

posterior rays shorter, forming a slightly convex

posterior margin to fin; longest rays extended as a

filament in large adults, length of longest ray 29.0-

33.2; 31.4-44.0.

Anal fin with 4 unbranched rays and 26-31,

usually 29, branched rays. Origin of anal fin behind midpoint of standard length 58.5-61.7; 57.6-

61.1. Fourth through eighth or ninth rays slightly

longer than posterior rays, forming a protruding

fin margin anteriorly and a straight margin posteriorly. Adult males with dorsally recurved or

straight spinules on posterolateral surface of posterior branches of anal-fin rays; these spinules

usually on third and fourth unbranched rays and

on first through sixth to twelfth branched rays.

Spinules occur from fin base to near fin tip in

first through fourth to seventh branched rays

(spinules sometimes sparsely scattered toward distal

ray tip), spinules on posterior successive rays cover

less area, usually present from anal-fin base to first

ray branch or to about midway between ray base

and first ray branch. One to several spinules per

bony ray segment. (Note: These are not hooks, but

spinules as described in Weitzman and Thomerson, 1970.)

Pectoral fin with 1 unbranched ray and 9-12,

usually 12, branched rays. Pectoral fins reach to

FIGURE 20.—Dentition of juvenile CarJana eigenmanni,

CAS(SU) 61380, 21.0 mm; Nicaragua, Rio Frio, about li/2

miles from San Carlos, Lake Nicaragua, collected 12 February

1963 by G. Myers.

FIGURE 21 Dentition of adult Carlana eigenmanni, USNM

74240, male, 56.4 mm; Costa Rica, La Junta, collected 8 April

1912 by S. Meek.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

or just anterior to pelvic-fin origin. Distance from

tip of snout to dorsal end of pectoral-fin base 26.9-

29.8; 24.4-26.6; length of pectoral fin from base

to tip of longest ray 17.8-21.3; 19.6-21.2.

Pelvic fin with 1 unbranched ray and 7-8, usually

7, branched rays, distal tip reaching slightly beyond anal-fin origin. Distance from tip of snout

to pelvic-fin origin 45.3-48.5; 41.6-45.9; pelvic-fin

length 16.7-19.3; 16.7-19.8. First or second through

fifth pelvic-fin rays with small spinules (occasionally only on first branched ray); 1 or 2 spinules

per bony ray segment.

Caudal fin with 10/9 principal rays (9/8 in 1

specimen); fin forked, not split to base. Caudal

fin symmetrical, without hooks or spinules. Dorsal

and ventral procurrent caudal rays equal in size.

Precaudal vertebrae 18-19, usually 18, total vertebrae 36-38, usually 37.

Color in alcohol: Ground color light yellowish

brown or pale brown. An extremely thin dark line

extends along midline. Melanophores under scale

margins form reticulate pattern dorsal to midline;

melanophores scattered over nape and head; below midline melanophores present to just above

anal-fin base and occasionally along myomere junctions between midline and anal-fin base. Melanophores usually less numerous in smaller specimens.

Adults with a series of about 8-10 vaguely defined

longitudinal bars along body, formed by patches

of larger melanophores under scales. Young with

2 small diffuse humeral spots composed of large

melanophores; adults with 1 humeral spot under

5 or 6 scales present in area of eighth or ninth

lateral scales, posterior to faint "pseudotympanum." Young with a dark blotch on caudal

peduncle, extending slightly onto caudal fin; adults

with a bar along midline from just anterior to a

vertical from adipose fin, widening at end of caudal

peduncle, then extending as a thin line to distal

tip of middle caudal-fin rays. Small melanophores

scattered on interradial membrane of all fins,

often concentrated along fin rays, not on them,

except in middle caudal-fin rays. Small specimens

with cheek silvery; peritoneum silvery except where

intestine is visible, forming an inverted dark gray

"comma" mark.

Range: According to Miller (1966; fide Bussing

and Rivas), C. eigenmanni is found from Lake

Nicaragua to the Atlantic slope of Costa Rica.

The type locality is La Junta, on the Rio Reventazon at an altitude of approximately 65 meters

(Meek, 1914).

COMPARISONS.—The generic placement of this

species has been discussed above. Carlana eigenmanni can be immediately distinguished from all

other Central American characids with a single

premaxillary tooth series by its number (26-31)

of branched anal-fin rays (see Table 3), shape of

the dentary teeth, and from all except Phenagoniates macrolepis by the presence of a fully toothed

maxillary in adults. See Figures 17-21 for illustration of changes occurring during growth.

Phenagoniates Eigenmann and Wilson

Phenagoniates Eigenmann and Wilson, 1914, in Eigenmann,

Henn, and Wilson, 1914:2 [type-species Phenagoniates wilsoni by monotypy].

Phanagoniates Eigenmann, 1915:43 [invalid spelling emendation].

Monotypic, separated from other Central American cheirodontin genera by an elongate body, large

tricuspid teeth, a long anal fin (42—50 branched

rays, all others treated here with 31 or fewer),

origin of anal fin anterior to dorsal-fin origin, low

number of pelvic-fin rays (i,5 versus i,7 in all

others), and no adipose fin.

A search of the literature indicates that Phenagoniates may be related to Leptagoniates Boulenger, differing only in the latter's possession of a

small adipose fin and possession of a complete

lateral line. We have examined Xenagoniates Myers

and found it to possess an adipose fin and ectopterygoid teeth. We also examined Paragoniates, which

also may be related to Phenagoniates, but which

has an adipose fin, is stouter, and has a long,

slender fully toothed maxillary. Reasonable estimation of relationships of these genera would

entail an investigation beyond the scope of this

paper. We do not consider Phenagoniates close to

other genera discussed in this paper and believe

that it lends credence to our conviction that the

Cheirodontinae is not a natural group.

Phenagoniates macrolepis (Meek and Hildebrand)

FIGURES 22, 23

Roeboides macrolepis Meek and Hildebrand, 1913:84 [original

description; Rio Cupe, Boca de Cupe, Panama].—Grey,

1947:183 [type listed], fig. 53 [holotype figured].
NUMBER 172

Phenagoniates wilsoni Eigenmann in Eigenmann, Henn, and

Wilson, 1914:2 [original description; Manigru].

Phanagoniates wilsoni.—Eigenmann, 1915:43 [description;

spelling emended], pi. V: fig. 1.

Phanagoniates macrolepis Meek and Hildebrand, 1916:272

[description; P. wilsoni considered a synonym]. Eigenmann, 1922:128 [listed from Atrato and Tuyra basins].—

Breder, 1927:177 [listed; variation; range], 163 [in key]

Jordan, Evermann, and Clark, 1930:95 [listed].—Hildebrand, 1938:249 [range].

Phenagoniates macrolepis Myers, 1942:90 [compared with

Xenagoniates bondi; note on spelling of Phenagoniates].—

Schultz, 1944:311 [listed from several localities in Venezuela]—Miller, 1966:784 [listed; range].

MATERIAL EXAMINED

•FMNH 7590, holotype, 45.1 mm, Panama: Rio Cupe, Boca

de Cupe, Darien, S. E. Meek and S. F. Hildebrand, 12 Feb.

1912.

•USNM 78665, Panama: Rio Cupe, Boca de Cupe, Darien,

S. E. Meek and S. F. Hildebrand, 24 Feb. 1912; 3 specimens

34.9-41.0 mm.

USNM 78661, Panama: Rio Cupe, Cituro, Darien, S. E. Meek

and S. F. Hildebrand, 25 Feb. 1912; 1 specimen 25.9 mm

(condition poor).

•USNM 78662, Panama, Rio Yape, Darien, S. E. Mark and

S. F. Hildebrand, 6 Mar. 1912; 1 specimen 42.4 mm.

•USNM 78664, Panama: Rio Aruza, Darien, S. E. Meek and

S. F. Hildebrand, 26 Feb. 1912; 1 specimen 39.7 mm.

•USNM 208517, Panama: Rio Sambu, Sta. BNW-20, approx.

7°52'N, 78°O5'W, 25 Mar. 1967; 4 specimens, all damaged,

approximately 30.0-43.0 mm (1 cleared and stained).

•USNM 208547, Panama: Rio Membrillo, Sta. BNW-9,

approx. 8°41'N, 77°41'W, 22 Mar. 1967; 3 specimens 41.4-

42.1 mm (1 damaged and unmeasurable).

•USNM 208546, Panama: Rio Morte at Hydro Station, Sta.

BNW-8, approx. 8°54'N, 77°33'W, 16 Mar. 1967; 3 specimens 33.4-33.6 mm (1 damaged and unmeasurable).

•FMNH 56541, Colombia: Certegui, C. Wilson, 1913; 1 specimen 23.8 mm (paratype of P. wilsoni).

31

•FMNH 56542 Colombia: Truando, C. Wilson, 1913; 1 specimen 30.3 mm (paratype of P. wilsoni).

We have examined paratypes of Phenagoniates

wilsoni and agree with previous authors that P.

wilsoni is synonymous with P. macrolepis.

Grey (1947) listed 5 paratypes of P. macrolepis;

this is an error as no paratypes were designated

in the original description. In one lot of P. macrolepis (USNM 78661) there are 2 fishes; 1 of these

is P. macrolepis, and the other specimen is in poor

condition and not identified here. This small fish

is a "cheirodontin," with a single row of elongate

conical teeth on the premaxillary and another

similar row along the entire anterior border of the

maxillary. The dentary teeth are elongate, with a

large median cusp and 2 smaller cusps, 1 on each

side. The anal-fin base is short and the ventral

procurrent caudal-fin rays are elongate. The specimen is too damaged for an accurate identification

or description.

Specimens used in description are included in

tables.

DESCRIPTION.—Standard length of specimens examined 33.4—45.1 mm. Body elongate, compressed

laterally; greatest body depth 26.0-29.8. Predorsal

body profile somewhat convex; profile arches dorsally just anterior to dorsal-fin base. Body profile

straight or very slightly convex between posterior

dorsal-fin base termination and procurrent caudalfin rays; adipose fin absent. Distance from eye to

dorsal-fin origin 44.8-47.0; distance from dorsal-fin

origin to end of caudal peduncle 45.1-49.1. Ventral

body profile gently rounded from jaws to anus;

steepest inclination ventral to jaws. Ventral body

profile protrudes ventrally its greatest distance just

FIGURE 22. Phenagoniates macrolepis, USNM 78665, 41.0 mm; Panama, Rio Cupe, Boca de Cupe,

Darien, collected 24 February 1912 by S. Meek and S. Hildebrand.
32

anterior to or just behind pelvic-fin insertion. Body

profile along anal-fin base straight or very slightly

convex; between posterior anal-fin termination and

procurrent caudal-fin rays, body profile concave.

Caudal-peduncle depth 8.7-10.2, peduncle length

5.8-7.1.

Head length 19.5-21.8. Eye diameter 8.0-9.5.

Snout length 4.6-4.9. Least bony interorbital width

5.6-6.2. Maxillary sloping ventrally and posteriorly,

forming an angle of 70-80 degrees to longitudinal

body axis; upper-jaw length 6.3-6.9. All teeth

tricuspid (except for smallest dentary and maxillary teeth, which are conical), with median cusp

larger; all teeth in a single series. Premaxillary,

anterior maxillary, and dentary teeth arise perpendicular to jaw bones and then curve posteriorly

just proximal to cusps. Maxillary with 8-11, usually

10, teeth; premaxillary with 6-7, usually 7, teeth

(holotype with 7); dentary with 10-12 teeth. No

teeth present on vomer, palatines, or pterygoids.

Fontanel moderate, length of that part anterior

to epiphyseal bar just over one-third length of that

part posterior to bar. Gill rakers short, 12-14 (holotype not counted). Circumorbital bones well ossified, infraorbital 3 wide, contacting preopercle

ventrally and posteriorly.

Scales moderately large, cycloid with concentric

FIGURE 23—Dentition of Phenagoniates macrolepis, USNM

208517, 41.5 mm; Panama, Rio Sambu, collected 25 March

1967 by BNW.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

circuli, and about 2-6 radii on exposed posterior

field. Lateral line with a slight ventral curve, incomplete, with 10-15 perforated scales. Lateral

scales about 41^42 (holotype damaged); scales

above lateral line 6-7, usually 7; scales below lateral line 6-7. Predorsal scales about 20—22. Scale

sheath along entire base of anal fin. Large axillary

scale present dorsal to pelvic-fin insertion; caudal

fin scaled at base.

Dorsal fin with 2 unbranched rays and 7-8

branched rays. Dorsal-fin origin posterior to analfin origin, about equidistant from eye and caudal

base. Distance from tip of snout to dorsal-fin origin

55.4-59.3. Third, fourth, or fifth dorsal-fin ray

longest with rays posterior to fifth abruptly shorter;

length of longest ray 22.8-25.4.

Anal fin with 4 unbranched rays and 45-50,

usually 49, branched rays. First unbranched ray

not always visible externally. Origin well anterior

to midpoint of standard length 41.5-45.5. Fourth

through eighth or ninth anal-fin rays slightly longer

than successive posterior rays, forming a slightly

protruding margin anteriorly and a straight margin

posteriorly. No anal-ray hooks present.

Pectoral fin with 1 unbranched ray and 11-12,

usually 11, branched rays. Pectoral fins reach well

beyond pelvic-fin insertion, almost to anal-fin

origin. Distance from tip of snout to dorsal end

of pectoral-fin base 20.2-22.4 and length of pectoral

fin from base to tip of longest ray 17.1—21.7.

Pelvic fin with i,5 rays in all specimens, distal

tip reaching just to anal-fin origin. Distance from

tip of snout to pelvic-fin insertion 33.0-34.8; pelvicfin length 8.8-10.2. No pelvic-ray hooks present.

Caudal fin with 10/9 principal rays in all specimens; fin forked, not split to base. Caudal-fin lobes

symmetrical. No hooks on caudal-fin rays. Dorsal

and ventral procurrent caudal rays usually of equal

size.

Precaudal vertebrae 13-14, usually 13. Total vertebrae 41-43, usually 42.

Color in alcohol: Ground color pale brown,

nape dark brown. Cheeks with few melanophores;

tip of lower jaw often dark brown. Back and sides

above midline with reticulate pattern of small

melanophores. Broad stripe of small melanophores

present from just behind opercle to caudal peduncle, extending to tips of middle caudal-fin rays;

ventral to stripe, melanophores sparsely distributed

along belly and sides, more concentrated posteriorly
NUMBER 172

(vaguely reticulate pattern anteriorly, following

myomere junctions above anal fin). Dorsal fin with

a lew melanophores along rays and with melanophores scattered on interradial membranes, more

concentrated distally. Caudal fin with melanophores

on borders of rays and on interradial membranes,

more concentrated on middle caudal rays. Anal fin

with melanophores on interradial membranes. Pectoral and pelvic fins with a few melanophores on

borders of fin rays.

Range: Phenagoniales macrolepis is found in

Central America only in eastern Panama, in the

drainages of the Rio Sambu, Rio Tuira, and Rio

Chucunaque (a major tributary of the Rio Tuira),

in Darien Province. The species was found by

Eigenmann (1915) in the Atrato basin in Colombia. Phenagoniates macrolepis is actually a northern South American fish which has invaded the

extreme eastern drainages of Panama; it does not

33

seem common, judging from collections we have

examined (this could simply be due to insufficient

sampling).

COMPARISONS.—Phenagoniates macrolepis is a distinctive species which cannot be confused with any

other recorded Central American cheirodontin. Its

long anal fin, dorsal-fin position, and large tricuspid teeth set it clearly apart from the other

species discussed in this work.

Saccoderma Schultz

FIGURES 24, 25, 26

Saccoderma Schultz, 1944:314 [type-species Saccoderma melanostigma Schultz, 1944, by original designation].

We have found only 1 collection of Saccoderma

species from Central America. There are 15 specimens of Saccoderma in SU 48851 (Panama: VeraFIGURE 24 Saccoderma hastata, holotype, FMNH 56383, male, 31.0 mm; Colombia, Soplaviento,

collected by C. Eigenmann (no date given).

guas Prov. vicinity of Santiago. Coll. T. D. White,

2 March 1956), along with 7 specimens of Cheirodon dialcpturus. Caudal squamation and dentary

teeth place these 15 specimens in Saccoderma, but

since they are immature we were unable to establish their specific identity.

We illustrate the caudal-fin squamation (Figure

26) of S. hastata (FMNH 56383, Colombia, Soplaviento), as an aid for future identification of the

genus from Panama. (Note that the lower [ventral]

caudal-fin rays of the holotype of S. hastata are

somewhat malformed and swollen; this is not true

FIGURE 2">.—Dentition of Saccodcrma liastaln, paratype,

FMNH 5G3OO, male, 26.9 mm; Colombia, Bernal Creek, collected by C. Eigenmann (no date given).
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of other specimens examined). The dentary teeth

(Figure 25) and modified saclike caudal-fin scales

(Figure 26) place Saccoderma clearly apart from

other genera included in this paper. In addition,

adult males of at least 3 species in the genus have

antrorse hooks on the rays of the lower caudal-fin

lobe.

Loftin (1965) states that he found Saccoderma

species in Panama on the Atlantic slope in the

Chagres basin and on the Pacific slope from the

Rio Bayano basin and coastal streams near the

Canal Zone. We were unable to find Saccoderma

in Loftin's Panama material at the ANSP or the

National Museum of Natural History.

Sumario

Las especies conocidas de peces quirodontinos

de la am6rica central, caracinos con solo una fila

de dientes premaxilaries, estan revisados. La clave

para su identification esta procedida. La revista

incluye miembros de los generos Cheirodon, Carlana, Phenagoniates, y Saccoderma. Todas estas

especies estan describidas y estan illustradas con

exception de la forma de Saccoderma de la america

central que solo se conoce en la forma juvenil y

puede ser que pertenezcan a una especie indescribable. Dos neuvas especies de Cheirodon estan

describidas, C. dialepturns de las cuencas pacificas

del oeste central y oeste de Panama, incluyendo la

FIGURE 2<5.—Caudal fin scalation of Saccoderma hastata, holotype, FMNH 56383, male, 31.0 mm;

Colombia, Soplavicnto, collected by C. Eigcnmann (no date given).
NUMBER 172 $5

parte de la Provincia Code oeste, hasta al Rio y Compsura estan sinonimozados con Cheirodon

Coto, Costa Rica, y C. mitopterus del Rio Code del basado en una reexaminacion de sujetos usados

Norte, una cuenca atlantica de Panama central. antes para definir estos generos asi como una nueva

Rhoadsia eigenmanni se encuentra en Carlana, evaluation del concepto del genero de peces caracel nombre generico disponible por el ya utilizado inos. Finalmente, unas razones estan presentadas

Carlia. Los generos Odontostilbe, Pseudocheirodon, para considerar los Cheirodontinae polyphyl^tico.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Branched dorsal-fin rays

Species

C. dialepturus

C. mitopterus

£. qorqonae

C. affinis

C. terrabae

C. eigenmanni

P. macro!epis

N 7

30

30

31

30

30

21

16 ]_

8

3

1

9

9

27

30

31

29

30

20

10 x

8.9

9.0

9.0

1 9.0

9.0

8.9

7.1

TABLE 2.—Branched pectoral-fin rays

Species

C. dialepturus

C. mitopterus

C^. gorgonae*

C. affinis

C. terrabae

C. eigenmanni

P. macrolepis

N

30

29

30

30

28

21

16

9

1

6

1

10

24

21

11

5

11

5

26

3

16

23

5

12

12

3

2

5

10

4

13 14 x

10.1

11.1

9.9

1 10.8

11.2

11.1

11.2

Holotype damaged

TABLE 3.—Branched anal-fin rays

Species N 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31/ 45 46 47 48 49 50 x

C. dialepturus

C_. mi topterus

C_. qorgonae

C. affinis

C_. terrabae

C_. eigenmanni

P. macrolepis

30 2 13 10 5

28 6 19 3

29 1 10 12 5 1

29 1 1 11 1_1

30 1

21

16

4 1

4 14 11

4 10 5 1

17.6

17.9

16.8

19.6

22.1

29.0

111 3 8 2 48.4
NUMBER 172 37

TABLE

Species

C. dialepturus

C. mitopterus

C. gorgonae

C. affini s

C. terrabae

C. eigenmanni

P. macrolepis

4 -

N

30

30

21

26

22

17

10

-Scales

5

26

21

12

above

6

30

4

14

21

2

lateral line

7 8 9

1

16 1

8_

X

6.0

5.1

5.0

5.5

6.0

8.1

6.8

Species

C. dialepturus

C. mitopterus

C. qorqonae

C. affinis

C. terrabae

C. eigenmanni

P. macrolepis

TABLE

N

30

30

21

29

24

18

12

5.—Scales

3 4

30

1 29

21

2

below

5

27

13

lateral line

6 7

11

12

6 ±

8 x

4.0

4.0

4.0

4.9

5.2

6 7.3

7.1

Species

C. dialepturus

C. mitopterus

C. gorgonae*

C.. affinis

C. terrabae

C. eigenmanni

P. macrolepis^

N 12

25

28

17

28

28

17

7 3

TABLE

13 14

1

2 2

6.—Gill

15 16

2

5 7

rakers

17

2

8

3

18

1

1

5

13

1

19

10

19

6

_6

5

20

9

5

2

1

7

21 22

5

3

15 1

X

19.7

19.4

18.2

18.0

20.4

15.9

12.8

* Holotype and juveniles not counted.

®Holotype not counted.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Species

C. dialepturus

C. mitopterus

C. gorgonae

C. affinis

C. terrabae

C. eigenmanni

P. macrolepis

N

30

00 O

CM CO 30

30

21

15

TABLE

1 2

2

22

1 26

10

7.—Maxillary

3

28

6

3

19

1

1

4

11

17

5

11

1

teeth

6 7

1

4 1

8 9

2 1_

1 3

10

1

9.

11 X

2.9

2.2

2.0

3.4

5.4

4.5

2 9.8

TABLE 8.—Total vertebrae

Species

C. dialepturus

C. mitopterus

C. qorgonae

C_. af finis

C. terrabae

C. eigenmanni

P. macrolepis

N 31

29

29

32

30

27

21

16

32

3

3

10

33

20

22

20

22

34

6

2

7

5

35 36

16 10

3

37 38/ 41

1

17 1

1

42 43 x

33.1

35.4

33.0

32.7

33.2

36.9

11 4 42.2
NUMBER 172 39

TABLE 9.—Morphometrics

Characters

Standard length

Greatest depth

Snout-Dorsal fin origin

Snout-Pectoral fin origin

Snout-Pelvic fin origin

Snout-Anal fin origin

Peduncle depth

Peduncle length

Pectoral fin length

Pelvic fin length

Dorsal fin length

Head length

Eye diameter

Snout length

Interorbital width

Upper jaw length

Eye-Dorsal fin origin

Dorsal fin originCaudal fin origin

Males

ranqe

25.6-32.6

32.2-36.2

50.5-54.3

23.8-26.8

44.3-47.5

62.1-66.4

12.6-15.0

14.5-16.1

20.8-26.2

17.9-24.2

29.9-33.8

23.5-25.3

8.6-10.2

4.8- 6.1

7.0- 8.0

7.6- 9.4

38.0-40.7

51.3-54.4

of C. dialepturus

Females

ranqe

25.4-33.3

33.7-38.3

52.1-55.2

23.4-26.7

43.7-48.8

62.3-66.2

12.1-13.8

13.9-16.5

22.1-24.2

18.5-20.6

27.3-32.3

23.3-24.6

8.8-10.5

4.7- 5.6

7.3- 8.0

7.1- 8.4

38.2-41.2

51.5-54.7

Holotype

28.0

35.0

54.3

26.8

47.5

66.4

14.3

15.0

23.9

18.2

*

25.1

9.3

5.0

7.5

8.6

40.7

50.0

* Damaged
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 10.—Morphometrics of C. mitopterus

Characters

Standard length

Greatest depth

Snout-Dorsal fin origin

Snout-Pectoral fin origin

Snout-Pelvic fi n origin

Snout-Anal fin origin

Peduncle depth

Peduncle length

Pectoral fin length

Pelvic fin length

Dorsal fin length

Head length

Eye diameter

Snout length

Interorbital width

Upper jaw length

Eye-Dorsal fin origin

Dorsal fin originCaudal fin origin

Males and

Females

range

24.9-36.8

23.6-38.3

40.6-68.4

18.7-30.2

34.9-60.0

49.7-85.0

8.5-14.7

13.1-21.0

15.0-32.5

15.3-30.1

23.2-39.8

18.5-28.7

7.9-12.9

4.3- 7.4

5.7- 9.2

6.2- 9.9

29.8-49.7

44.0-74.3

Holotype

35.0

34.3

59.8

27.3

53.5

75.2

14.3

20.6

32.5

30.1

•

26.6

12.2

6.6

8.4

9.4

44.8

68.2

Damaged
NUMBER 172 41

TABLE 11.—Morphometrics of C. gorgonae

Characters

Standard length

Greatest depth

Snout-Dorsal fin origin

Snout-Pectoral fin origin

Snout-Pelvic fin origin

Snout-Anal fin origin

Peduncle depth

Peduncle length

Pectoral fin length

Pelvic fin length

Dorsal fin length

Head Length

Eye diameter

Snout length

Interorbital width

Upper jaw length

Eye-Dorsal fin origin

Dorsal fin originMales

range

22.9-26.7

33.4-37.8

51.4-54.6

24.4-28.8

45.0-48.9

62.2-66.9

12.0-14.3

13.6-17.1

21.4-25.4

18.3-21.4

29.6-32.2

24.0-26.3

9.3-10.7

5.4- 5.8

6.5- 8.1

7.4- 8.0

38.0-41.5

51.2-53.7

Females

range

23.1-28.4

33.4-37.5

52.6-56.8

24.2-28.0

45.5-49.8

62.5-67.2

12.6-13.4

13.6-15.8

20.2-22.1

16.5-18.7

27.6-32.4

23.5-26.0

9.2-10.8

5.3- 5.8

7.4- 7.9

7.4- 7.9

38.7-43.2

50.2-54.5

Juveniles

19.4-21.3

25.7-33.5

49.5-52.0

26.7-28.6

46.9-49.0

63.1-65.0

11.1-12.9

14.1-15.1

21.8 *

16.0-18.0

29.8 *

25.3-27.3

10.8-11.9

5.4- 5.8

7.0- 8.3

7.6- 8.3

36.6-37.9

49.3-52.7

Holotype

22.1

30.3

53.9

28.5

48.0

65.6

11.8

14.0

*

*

*

25.3

10.4

5.4

7.7

8.1

39.8

49.4

Damaged
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 12.—Morphometries of C.

Characters

Standard length

Greatest depth

Snout-Dorsal fin origin

Snout-Pectoral fin origin

Snout-Pelvic fin origin

Snout-Anal fin origin

Peduncle depth

Peduncle length

Pectoral fin length

Pelvic fin length

Dorsal fin length

Head length

Eye diameter

Snout length

Interorbital width

Upper jaw length

Eye-Dorsal fin origin

Dorsal fin originCaudal fin origin

Males and

Females

range

22.5-38.8

37.0-44.3

51.7-54.5

24.5-27.1

46.1-50.4

63.1-68.3

12.4-14.1

11.4-15.5

20.1-23.6

17.0-20.2

30.0-34.5

23.2-26.3

9.0-10.6

5.3- 6.2

8.2- 9.2

6.0- 7.6

38.4-42.7

52.6-56.0

affirm

Hoiotype

29.5

39.0

54.0

24.8

47.5

64.8

12.5

14.2

20.6

16.9

30.5

24.4

10.2

5.8

8.5

6.4

39.0

53.6

TABLE 13.—Morphometncs

Characters

Standard length

Greatest depth

Snout-Dorsal fin origin

Snout-Pectoral fin origin

Snout-Pelvic fin origin

Snout-Anal fin origin

Peduncle depth

Peduncle length

Pectoral fin length

Pelvic fin length

Dorsal fin length

Head length

Eye diameter

Snout length

Interorbital width

Upper jaw length

Eye-Dorsal fin origin

Dorsal fin originCaudal fin origin

of C. terrabae

Males and Females

range

29.8-49.8

32.9-42.6

50.0-57.6

24.2-29.0

45.1-51.1

61.3-69.5

11.0-13.2

11.3-15.0

19.4-24.1

16.7-21.3

27.8-34.4

22.8-26.6

8.7-10.4

5.4- 6.5

7.6- 9.2

6.0- 7.7

37.2-43.0

51.0-58.0
NUMBER 172 43

TABLE 14.—Morphometrics of C.

Characters

Standard length

Greatest depth

Snout-Dorsal fin origin

Snout-Pectoral fin origin

Snout-Pelvic fin origin

Snout-Anal fin origin

Peduncle depth

Peduncle length

Pectoral fin length

Pelvic fin length

Dorsal fin length

Head length

Eye diameter

Snout length

Interorbital width

Upper jaw length

Eye-Dorsal fin origin

Dorsal fin originCaudal fin origin

Juveniles

ranqe

21.0-29.8

32.0-39.5

53.4-56.7

26.9-29.8

45.3-48.5

58.5-61.7

8.3-10.1

8.7-10.9

17.8-21.3

16.7-19.3

29.0-33.2

24.9-27.6

10.8-12.7

4.9- 6.9

7.2- 9.0

7.5- 8.2

38.9-41.5

49.0-52.4

eigenmanni

Adults

ranqe

34.3-56.4

35.8-41.2

52.0-55.0

24.4-26.6

41.6-45.9

57.6-61.1

9.5-11.6

9.5-10.5

19.6-21.2

16.7-19.8

31.4-44.0

23.1-25.2

8.5-10.8

6.1- 7.1

8.4- 8.9

8.6-11.9

39.6-41.2

51.9-56.3

Hoio type

52.8

41.2

53.1

24.6

42.7

59.4

11.4

10.0

20.8

19.5

36.4

23.6

9.1

6.6

8.9

11.6

40.7

55.8
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 15.—Morphometrics of P. macrolepis

Characters

Standard length

Greatest depth

Snout-Dorsal fin origin

Snout-Pectoral fin origin

Snout-Pelvic fin origin

Snout-Anal fin origin

Peduncle depth

Peduncle length

Pectoral fin length

Pelvic fin length

Dorsal fin length

Head length

Eye diameter

Snout length

Interorbital width

Upper jaw length

Eye-Dorsal fin origin

Dorsal fin originCaudal fin origin

Males and Females

range

33.4-45.1

26.0-29.8

55.4-59.3

20.2-22.4

32.1-34.8

41.5-45.5

8.7-10.2

5.8- 7.1

17.1-21.7

8.8-10.2

22.8-25.4

19.5-21.8

8.0- 9.5

4.6- 4.9

5.6- 6.2

6.3- 6.9

44.8-47.0

45.1-49.1

Hoiotype

45.1

29.0

56.8

20.8

32.1

41.7

10.2

7.1

18.0

8.9

23.5

19.5

8.0

4.9

6.0

6.9

46.8

48.1
NUMBER 172 45

Literature Cited

Astorqui, I.

1971. Peces de la cuenca de los grandes lagos de Nicaragua. Revisita de Biologica Tropical 19(l,2):7-57.

Bohlke, J. E.

1952. Studies on Fishes of the Family Characidae, No. 2:

The Identity of the South American Characid Fish

Genus Distoechus Gomez. Annals and Magazine of

Natural History, series 12, 5:793-795.

1954. Studies on the Phylogeny and Systematics of Fishes

of the Family Characidae. Unpublished doctoral

dissertation, Stanford University.

1958. Studies on Fishes of the Family Characidae, No. 16:

A New Hyphessobrycon from Costa Rica. Bulletin

Florida State Museum of Biological Sciences, 3(4):

173-178.

Breder, C. M., Jr.

1925. Notes on Fishes from Three Panama Localities:

Gatun Spillway, Rio Tapia and Caledonia Bay.

Zoologica (New York), 4:137-158.

1927. The Fishes of the Rio Chucunaque Drainage, Eastern Panama. Bulletin of the American Museum of

Natural History, 62:91-176.

Bussing, W. A.

1967. New Species and New Records of Costa Rican

Freshwater Fishes with a Tentative List of Species.

Revisita de Biologica Tropical, 14(2):205-249.

Cope, E. D.

1870. Contribution to the Ichthyology of the Maranon.

Proceedings of the American Philosophical Society,

11:559-570.

Eigenmann, C. H.

1915. The Cheirodontinae, a Subfamily of Minute Characid Fishes of South America. Memoirs of the

Carnegie Museum, 7(1): 1-99.

1922. The Fishes of Western South America, Part 1.

Memoirs of the Carnegie Museum, 9(1): 1-278.

Eigenmann, C. H., A. Henn, and C. Wilson

1914. New Fishes from Western Colombia, Ecuador, and

Peru. Indiana University Studies, (19): 1-15.

Eigenmann, C. H., and G. S. Myers

1929. The American Characidae. Memoirs of the Museum

of Comparative Zoology, 43(5):429-574.

Evans, H. E.

1952. Notes on Some Fishes from Central Panama and

the Canal Zone. Copeia, 1952(l):43-44.

Evermann, B. W., and E. L. Goldsborough

1909. Notes on Some Fshes from the Canal Zone. Proceedings of the Biological Society of Washington,

22:95-104.

Gery, J.

1965. A New Genus from Brazil—Brittanichthys. Tropical

Fish Hobbyist, 13(6): 13-23, 61-69.

1972. Contribution a L' Etude de Poissons Characoides de

I' Equateur. Ada Humboldtiana, Series Geologica,

Paleontologica el Biologica, 2:1-110, plates i-viii.

Girard, C. F.

1854. Abstract of a Report to Lieut. James M. Gillis,

U.S.N., upon the Fishes Collected during the U.S.

Naval Astronomical Expedition to Chile. Proceedings of the Academy of Natural Sciences, Philadelphia for 1854-1855, 7:197-199.

Gosse, J. P.

1966. Poissons d'eau douce du versant Pacifique du Costa

Rica et de Panama recoltes par sa majeste le roi

Leopold de Belgique. Bulletin lnstitut Royal des

Sciences Naturalles de Belgique, 42(28): 1-24.

Gray, J. E.

1845. Catalogue of the Specimens of Lizards in the Collection of the British Museum. 289 pages. London.

Grey, M.

1947. Catalogue of Type Specimens in Chicago Natural

History Museum. Fieldiana: Zoology, 32 (3): 109-205.

Hennig, W.

1966. Phylogenetic Systematics. 263 pages, 69 figures,

Urbana: University of Illinois.

Hildebrand, S. F.

1938. A New Catalogue of the Freshwater Fishes of Panama. Field Museum of Natural History, Zoological

Series, 22(4): 219-359.

Jordan, D. S., B. W. Evermann, and H. W. Clark

1930. Check List of the Fishes and Fishlike Vertebrates

of North and Middle America North of the Northern Boundary of Venezuela and Colombia. Report

of the United States Commission of Fisheries for

1928, (2): 1-670.

Loftin, H. G.

1965. The Geographical Distribution of Freshwater Fishes

in Panama, viii + 264 pages, 1 map. Unpublished

doctoral dissertation, Florida State University.

L6pez, M. I.

1972. Contribucion al estudio de los peces Cheirodontinae

(Familia Characidae) de Costa Rica y Panama:

Generos Compsura y Pseudocheirodon. Revisita de

Biologica Tropical, 20(l):93-129.

Meek, S. E.

1912. New Species of Fishes from Costa Rica. Field Museum of Natural History, Zoological Series, 10(7):

69-75.

1914. An Annotated List of Fishes Known to Occur in

the Freshwaters of Costa Rica. Field Mttseum of

Natural History, Zoological Series, 10(10): 101-134.

Meek, S. E., and S. F. Hildebrand

1913. New Species of Fishes from Panama. Field Museum

of Natural History, Zoological Series, 10(8):77-91.

1916. The Fishes of the Freshwaters of Panama. Field

Museum of Natural History, Zoological Series,

10(15):2l7-374.

Miller, R. R.

1966. Geographical Distribution of Central American

Freshwater Fishes. Copeia, 1966(4):773-801.
46

Myers, G. S.

1942. Studies on South American Freshwater Fishes, I.

Stanford Ichthyological Bulletin, 2(4):89-114.

1958. Trends in the Evolution of Teleostean Fishes.

Stanford Ichthyological Bulletin, 7(3):27-30.

Nelson, K.

1964. Behavior and Morphology in the Glandulocaudine

Fishes (Ostariophysi, Characidae). University of

California Publications in Zoology, 75(2):59-152.

Schultz, L. P.

1944. The Fishes of the Family Characinidae from VeneSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

zuela, with Descriptions of Seventeen New Forms.

Proceedings of the United States National Museum,

95:235-367.

Strand, E.

1926. Miscellanea nomenclatorica zoologica et paleontologica, I—II. Archiv fur Naturgeschichte, 1926, part

A, 8:30-75.

Weitzman, S. H., and J. E. Thomerson

1970. A New Species of Glandulocaudine Characid Fish,

Hysteronotus myersi, from Peru. Proceedings of the

California Academy of Sciences, 38 (8): 139-156.

11/15


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