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The characiform fishes of the Apure River drainage, Venezuela

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Title:
The characiform fishes of the Apure River drainage, Venezuela
Creator:
Taphorn, Donald C., 1951-
Publication Date:
Language:
English
Physical Description:
2 v. (vi, 892 leaves) : ill. ; 29 cm.

Subjects

Subjects / Keywords:
Animal fins ( jstor )
Couplets ( jstor )
Fish ( jstor )
Fish scales ( jstor )
Food ( jstor )
Natural history ( jstor )
Pigmentation ( jstor )
Rivers ( jstor )
Species ( jstor )
Teeth ( jstor )
Characiformes ( lcsh )
Genre:
bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1990.
Bibliography:
Includes bibliographical references (leaves 864-886).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Donald C. Taphorn.

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University of Florida
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University of Florida
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Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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AA00004770_00001 ( sobekcm )

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THE CHARACIFORM FISHES
OF THE APURE RIVER DRAINAGE, VENEZUELA

















By

DONALD C. TAPHORN


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA


1990












ACKNOWLEDGMENTS


Saying thank you is hardly adequate to repay the many people who have

helped me in this project. They have given me tremendous gifts: precious

time, sound advice and useful suggestions, as well as moral and financial

support and many other things. They helped get the fish out of the water

and into a jar in the museum, get the sometimes jumbled ideas in my head

into some semblance of order, get my body out of bed and into the lab, and

get the words out of the computer and onto this page. But thank them I

must and I do so sincerely.

My parents and family are first in this list of special people who

have helped. They are always with me in spite of the large distances that

have separated us. Next I thank the many friends, professors, colleagues

and students who have helped to make this dissertation possible. They are

Dr. Carter Gilbert, cochairman of my committee for these many years and

tireless in the dreary task of wading through and correcting the rough

drafts, Drs. Horst Schwassmann, Frank Nordlie, and Nigel Smith, who pa-

tiently served on my committee and suggested many helpful improvements and

Dr. Craig Lilyestrom, his wife Maria and their children Lynda and Wayne who

shared a home with me in Venezuela. During my visits to Gainesville, I

enjoyed the warm hospitality of Dr. and Mrs. Carter Gilbert, Mr. and Mrs.

George Burgess, and Mr. and Mrs. Terry Converse.

Dr. Kirk Winemiller, Mr. Leo Nico and Mr. Stewart Reid worked on

postgraduate degrees in our laboratory, shared in the field work, and






generously provided specimens and information on the ecology of Apure drain-

age fishes from their master's and doctoral work.

I also thank the friends and colleagues who helped pull the other

(usually the deep) end of the seine, Craig Lilyestrom, Craig and Carmen

Olds, Leo Nico, Stewart Reid, Kirk Winemiller, Oscar Leon M., Naboth

Montilla A., Mike Taphorn, Pablo Osman, Larry Page, Carter Gilbert,

Richard Franz, Jack Karr, Jamie Thomerson, Guillermo and Ram6n Feo,

Alex Flecker and many others too numerous to mention here. Special

thanks go to Ing. RNR Angelina Licata and Mr. Oscar Leon Mata for the

drawings and to Lic. Jairo Perez and Ing. Aniello Barbarino D. for the

maps. Three technicians have worked with me in the Fish Collection,

Aldo Garcia, Eric Sutton and Keyla Marchetto; I am greatly indebted to

them for their assistance with the unending museum chores of processing

and cataloging the specimens, preparing labels and card files, and

keeping the computer records up to date. Several of my students in the

Environmental Engineering Department have done thesis work on fishes,

and many also held student jobs in the Fish Collection and helped with

the museum chores. Special thanks go to Pablo Osman, Norberto Saavedra,

Enrique Marzola, Aniello Barbarino Duque, Cecelia G6mez, Guillermo

Cedefo, Ricardo Smith, Yury HernAndez. I also thank Dr. Richard Schar-

gel for his helpful talks about Apure drainage soils and Prof. Heberto

Pacheco and the other personnel of the Cartographic Center of UNELLEZ

who helped obtain topographical maps.

The Secretario de Investigaci6n de la UNELLEZ, FONAIAP, FUNDA-

CITE, CONICIT all provided financial support for this project.












TABLE OF CONTENTS


page


ACKNOWLEDGMENTS.............


........ i i


ABSTRACT..............................................
INTRODUCTION.....................................


MATERIALS AND METHODS.......
Collecting...............
Curation.................
Material Examined........
Format...................
RESULTS.................
Characiformes...........
Anostomidae.............
Characidae ..............


Characidiidae ....................


Chilodontidae.....
Ctenoluciidae.....
Curimatidae.......
Cynodontidae......
Erythrinidae......
Gasteropelecidae..
Hemiodontidae.....
Lebiasinidae......
Parodontidae......
Prochilodontidae..


DISCUSSION..........................................
Ecological Considerations .......... ...............
Zoogeographical Considerations....................


21
23
31
100
606
664
670
677
721
739
760
766
777
796
808

823
823
854


LITERATURE CITED..................................... 864

APPENDIX.............................................. 887
BIOGRAPHICAL SKETCH.................................. 891


11111


Ilfllll


11111.11


11111.11













Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

THE CHARACIFORM FISHES
OF THE APURE RIVER DRAINAGE, VENEZUELA

By

DONALD C. TAPHORN

May, 1990

Chairman: Dr. Carter R. Gilbert
Major Department: Zoology

The Apure River drainage of Venezuela collects waters from many

different aquatic habitats that vary greatly in geology, climate, vegeta-

tion and altitude. Over ten years of sampling has yielded 354 species of

freshwater fishes pertaining to ten orders and forty-one families. This

report includes taxonomic synonymies, keys, illustrations, morphological

descriptions, distribution maps, data on abundance, habitat, life history

strategy and diet for 138 Characiform species of the families Anostomidae,

Characidae, Characidiidae, Chilodontidae, Ctenoluciidae, Curimatidae,

Cynodontidae, Erythrinidae, Gasteropelecidae, Hemiodontidae, Lebiasinidae,

Parodontidae, and Prochilodontidae.

Fish distribution within the Apure River drainage is primarily gov-

erned by two parameters, altitude and water chemistry. The few characi-

forms adapted to mountain and piedmont conditions tend to be restricted

in distribution to only those areas.






In the llanos, two assemblages are recognized. Whitewater, muddy

streams are the most common habitat in the drainage, but few characiforms

are restricted only to them. Blackwater habitats with distinct fish assem-

blages exist in isolated pockets in the Aguaro-Guariquito drainage, in the

Caicara Creek drainage of northern Apure state, and in west-central Barinas

state.

The drastic changes in climate between wet and dry season in the

Apure River drainage have molded characiform life history strategies. Most

characiforms have adopted an rl or r2 life history strategy. Of 68 species

that have adopted the rl strategy, most are small omnivorous fishes with

low juvenile and adult survivorship, low fecundity per reproductive bout

(but repeated reproduction throughout the rainy season), short generation

times (many live less than one year), and widely fluctuating population

densities during the year. The other 70 species present have been classi-

fied as r2 strategists. These species typically are larger, are often

migratory, and reproduce annually in a single massive spawning event.

Juvenile survivorship is low, but adults live for several years and reach

larger sizes. Populations fluctuate greatly from wet to dry season because

of the high juvenile mortality. No characiforms in the Apure drainage have

adapted the K-strategy.

The Apure drainage ichthyofauna is a mosaic of relatively young

species that have evolved in the whitewater habitats and blackwater species

probably derived from an older Guyana Shield fish fauna.













INTRODUCTION


Tropical aquatic ecosystems are suffering rapid modification and

deterioration as man's growing population makes ever-increasing demands for

water, forest products, and agricultural land. Many Venezuelan rivers are

already so polluted that we will never know much, if anything, about their

original fish faunas. Fishes are suffering loss of habitat to wetlands

drainage programs, habitat modification by dams and dikes, pollution from

urban sewage, agricultural fertilizers and biocides sedimentation from

deforestation and poor upland watershed management, overfishing, and the

introduction of exotic species.

Efforts to produce effective management plans for a rational develop-

ment of tropical aquatic resources suffer from an almost total lack of basic

information about the aquatic organisms that occupy these systems. Not even

lists of the fishes, mollusks, crustaceans, aquatic insects, plants, etc.,

can be found for most tropical rivers, much less data on abundance, life

histories, critical habitat requirements or other aspects of their biology.

Furthermore, the majority of these organisms have never been sufficiently

sampled, nor has their taxonomy been adequately studied. The literature

provides few guides to the identification, and for the most part, original

species descriptions are hidden in old primary references not usually avail-

able to workers in South America. What little information is available is

seldom written in Spanish, and so is of little use to management officials.

For these and many other reasons, inventories of tropical aquatic organisms






are desperately needed. The decision to undertake a faunal survey of the

Apure River drainage, part of which forms the basis of my doctoral disserta-

tion, is in response to this need.

In South America the Orinoco River Basin (fig. 1) collects the waters

of a vast area covering approximately 1,123,000 km2, second only (although

the Rio La Plata is longer) to the immense Amazon River Basin to the south,

which drains some 7,000,000 km2 (Zinck 1977, Smith 1981). About three-

fourths of the area of the Orinoco Basin falls under Venezuelan jurisdic-

tion, whereas the rest is Colombian. The river traverses some 2,148 km in

its journey to the sea from its currently recognized source near Cerro

Delgado-Chalbaud in the ancient mountains of the Guyana Shield near the

border with Brazil. In my opinion, the true source of the Orinoco is the

Guaviare River, which originates in the Andes of Colombia and contributes

much more water than the Ventuari. In any case, the Orinoco flows west from

its Venezuelan source, at first descending through dense tropical forest

that grows in the well-weathered rocky drainages of the Guyana highlands.

After sharing a portion of its reddish-brown waters with the Amazon via the

Casiquiare Channel, it receives its first major tributary, the Ventuari, and

then veers north to run along the northwestern edge of the Guyana Shield

where it marks the boundary between Colombia and Venezuela. Several major

tributaries enter the Orinoco from Colombia in this stretch (fig. 1), among

the largest of which are the Guaviare, Vichada, Tomo and Meta. At Puerto

Piez, the Orinoco turns slightly to the east, and receives the black waters

of the Cinaruco and Capanaparo rivers that drain the grassland savannas of

southern Apure state. Rocks of the Guyana Shield still project though sandy

bottom sediments at this stage. Gradually, the river bends more eastward.

Just before reaching Caicara, the muddy waters of the Apure River dump



















































































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their Andean sediments into the main channel of the Orinoco. Since the

Orinoco is often full during peak flood of the Apure, it tends to hold back

the waters of the Apure, and has formed an interior delta where vast areas

flood each year. The Orinoco changes course at this point to run in a

northeasterly direction towards the sea. It is bordered by the semi-arid

central plains to the north, and the humid Guyana Shield to the south. It

picks up vast quantities of black waters from the Caura and Caroni rivers

before it enters its highly productive delta and flows into the Atlantic

Ocean just south of the island of Trinidad. The average annual outflow is

about 33,000 m3/s (Zinck 1977). The Amazon, by way of comparison, delivers

an annual average of about 175,000 m3/s (Smith 1981).

The Apure River drainage lies mostly within 7" and 10 North lati-

tudes, and 66" and 720 West longitudes. It is Venezuela's largest tributary

(in terms of drainage area) of the Orinoco (fig. 2a & 2b) and drains 167,000

km2, or nearly 15% of the total area of the Orinoco Basin. Its average flow

of nearly 2000 m3/s (Zinck 1977) divided by the drainage area gives a figure

of about 378 mm/year runoff, which is about 24% of the mean precipitation

(Saunders & Lewis 1988a). It is exceeded in Venezuela by only two other

tributaries, both blackwater rivers from the nutrient poor soils of the

Guyana Shield, where rainfall is much higher. The Caroni River, which

drains some 93,000 km2 delivered over 4,000 m3/s before the enormous Rail

Leone (Guri) Dam was built. The Caura River, slated for future hydroelec-

tric development, carries 2,700 m3/s. The Guaviare River which drains

159,500 km2 and the Meta River with 103,000 km2 are two of the major Orinoco

tributaries in Colombia (Zinck 1977). The headwaters of the Apure drain the

southeastern flank of the Andes, and are located approximately 800 km by

river from the Orinoco River. Over 15% of the drainage is mountainous






















































































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(above 500m in elevation). Transit time from the Andes to the Orinoco

varies seasonally in the range of one to four weeks (Saunders & Lewis

1988a,b). On the way, the numerous branches of anastomosing rivers cross

the llanos, flat alluvial plains that formed in the late Pleistocene over

older marine sediments. In the wet season, most of the lower part of the

drainage is inundated by sheet flooding. The floodplain of the Apure is

much more extensive than that of most rivers. An internal delta of about

5000 km2 forms where the Apure and the Arauca (the first major tributary of

the Orinoco to the south) flow into the Orinoco. When discharge is high in

the Orinoco, a large inland sea forms between the mouths of the Capanaparo

river (the next Orinoco tributary to the south) and the Apure. The alter-

nation of wet and dry seasons causes annual changes in river level of about

six to eight meters. Using river level values from San Fernando de Apure,

Saunders & Lewis (1988a) defined four hydrologic phases to describe the

relationship between the river and the floodplain. The river is not in

contact with the floodplain during the first, low-water phase, which corre-

sponds to stage heights below 38 m. The rains, which usually start in

April or May, initiate the second, rising-water phase, during which the

river is still within its banks (38-42m). The inundation phase commences

when stage heights surpass 42m, and river water floods onto the surrounding

floodplain. Flooding usually begins in July, and peaks in September or

October. When the dry season sets in and water levels drop to below 42m,

the fourth, or falling-water phase begins (42-38m).

Of all of the Orinoco tributaries, the Apure is by far the most acces-

sible, with good, all-season roads crossing all of its major drainage sys-

tems. This factor alone was enough to select it as my study site, but in

addition to easy access the climate and topography are extremely





11

diverse. As a result, many different aquatic biotopes are included in the

drainage. There are cold mountain streams that tear through the Andes

at breakneck speeds; tepid, blackwater lowland swamps; large "white-

water" muddy rivers that meander slowly through the lower llanos (a

Venezuelan term for plains or savanna); and all the varied intergrada-

tions of these that can be imagined. The good roads in this region owe

their existence to the fertile soils and resultant extensive agriculture

that has developed. Deforestation has proceeded virtually unchecked

since colonial times. As more and more forests give way to pastures and

crops, the aquatic fauna becomes increasingly degraded. Massive fish

kills occur almost as a matter of routine when the first rains fall

after the months of drought and wash the accumulated pesticides and

fertilizers into the drainage in one large dose. Natural aquatic habi-

tats are disappearing rapidly and fish populations are suffering accord-

ingly. This too gave the Apure drainage a high priority as a study

area, inasmuch as the fishes sensitive to these changes may not be

around indefinitely.

Another factor in the selection of the Apure drainage is that the

Apure River and its larger tributaries support a large commercial fishery,

and provide a significant source of protein and extra income for rural

families. Almost no basic biological data is available for most species.

Declining catches in recent years and local conflicts over the best use of

aquatic resources in the region has local and federal governments concerned.

Most commercial species are migratory, moving in tune with the drastic

seasonal changes in water level and flow because of the accentuated wet

and dry seasons typical of the western llanos. Funding for impact

studies of local dams on commercial species helped pay for the sampling.








The Andean piedmont was characterized as an area of recent speciation

by Mago L. (1978), but Andean piedmont rivers, flowing out through narrow

valleys into the plains, offer excellent opportunities for hydroelectric

power generation. The Venezuelan government, eager to diversify its energy

resources now that the end of their vast petroleum wealth is in sight, has

projects for dams on almost every major piedmont river system. These dams

have denied the commercial species, particularly the coporo, Prochilodus

maria, access to the upland piedmont and montane lotic systems, and disrupt

the annual cycle of flood and drought used by migratory species as the cue

to make their moves. Commercial species populations are on the decline

throughout the Apure drainage, but direct causes are not easily identified

because of the wide array of concurrent negative impacts that are occurring.

As the commercial fish catch drops, concern is mounting, and governmental

interest in finding a solution is growing. But again, lack of basic infor-

mation about fish populations and ecology hinders progress.

In his seminal work, "Lista de los Peces de Venezuela incluyendo un

Estudio Preliminar sobre la Ictiogeografia del Pais," Mago L4ccia (1970)

lamented the deplorable state of knowledge of the Venezuelan freshwater fish

fauna. He noted that most of the pioneering ichthyological works of Eigen-

mann, Steindachner, Regan, Kner, Pellegrin, Norman, Fowler and others were

based on specimens from just about everywhere in South America except Vene-

zuela. Only the Maracaibo Basin, studied in detail by Leonard P. Schultz

(1944a & b, 1949), was relatively well known. Since then, the pace of

ichthyological research in Venezuela has quickened considerably. This is

principally due to the work of Dr. Mago L. and his students, but contribu-

tions have also been made by Agustin Fernandez YUpez, Felipe Martin and

Manuel Ramirez. However, the present state of knowledge on the freshwater








fishes of Venezuela is far from complete, and most areas still need to be

better sampled.

Neotropical characiform fishes have received considerable attention

from ichthyologists. C. H. Eigenmann provided the basis for most modern

studies with a series of papers published in the late 1800's and early

1900's that culminated in several huge tomes such as his American Characidae

series. Other workers such as E. Ahl, J. B6hlke, H. Fowler, J. Gdry, J.

Hoedeman, R. von Ihering, W. Ladiges, K. Luling, S. Meek, H. Meinken, N.

Menezes, G. Myers, A. and R. Ribeiro de Miranda, T. Roberts, H. Travassos,

S. Weitzman and R. Vari have continued the effort to give us a much better

idea of the tremendous diversity of form and function represented in this


group.













MATERIALS AND METHODS


Collecting

Many different collecting methods were employed, but most collections

were made with small, fine-meshed seines. Gill nets (both multifiber nylon,

and monofilament), beach seines, cast nets (both fine and commercial mesh

sizes), hand nets, and small otter trawls (in the Apure River's main chan-

nel) were also used. We also fished with electrofishing gear, rotenone, and

hook and line. Some specimens were obtained from commercial fishermen.

Fishing effort was not vigorously controlled at each site. Usually

sampling continued until no new species appeared. From 1977 to 1989 we

made a total of 875 collections in the Apure drainage (fig. 2b).

Curation

Specimens were preserved in 10-15% formalin solution, and later trans-

ferred to either 50% isopropanol, or 70% ethanol. Larger specimens were

usually slit on the right side to ensure complete preservation.

Material Examined

Most of the specimens examined as part of this study have been depos-

ited in the Museo de Ciencias Naturales de la UNELLEZ--Guanare (MCNG).

Additional material is deposited in the Field Museum of Natural History in

Chicago (FMNH), the museum of the Illinois Natural History Survey (INHS),

the Museo de Biologfa de la Universidad Central de Venezuela in Caracas

(MBUCV), the museum of Sacramento State College, in Sacramento California

(SSC), the Texas Memorial Museum in Austin (TMM), the Florida Museum of






Natural History in Gainesville (UF), and the United States National Museum

(USNM) in Washington. Other museum acronyms used in this study are:

AMNH--American Museum of Natural History; ANSP--Academy of Natural Sciences,

Philadelphia; BMNH--British Museum of Natural History; CAS--California

Academy of Sciences; CM--Carnegie Museum (now mostly in FMNH and CAS);

IU--Indiana University (these specimens are now located mostly in FMNH or

CAS); MCZ--the Museum of Comparative Zoology at Harvard; MNHN--Musdum

National d'Histoire Naturelle, Paris; MZUSP--the Museum of Zoology of

the Sao Paulo University in Brazil; NMW--the Vienna Natural History

Museum; SU--Stanford University (now at CAS, and designated CAS-SU);

ZMA--Zoologisch Museum of Amsterdam; ZMB--Zoologisch Museum of Berlin.

Format

The following format has been used to organize the information pre-

sented here. A brief introduction summarizing pertinent data for each

family is followed by a key to the species known to be present in the Apure

drainage. These sections are followed by the species accounts, which are

arranged, in sequence, under the following headings:

1. Genus--species--author--year (centered at top).

2. Common Name in Spanish & Common Name in English (if known).

3. Fig. #. This is the figure numbers) of the illustrationss.

4. Map: fig. #. This is the figure number of the dot map showing the

Apure drainage distribution.

5. Couplet. #. The couplet(s) where the species keys out is given.

6. Generic synonymy. This section, given as part of the species account

for the first species of each genus, includes the principal elements of a

generic synonymy: original name, describer, date, and type species. Only

primary junior synonyms are listed.






7. Specific Synonvmy. The specific synonymy includes the original name,

describer, date, and page number, as well as the type locality (only given

for citations of original descriptions), junior synonyms, and selected

references to Venezuelan specimens or other important reports. It is not

intended as a complete synonymy, but in many cases it may be. In most cases

synonymies have been copied from the literature. When the original descrip-

tion has been seen by me (many are in unavailable literature) this is so

indicated (as: seen).

8. Types. If readily available, museum location of the type material is

presented.

9. Comments. Comments are given on the taxonomic status of the species.

A synopsis of recent opinions on the status of the species or an explanation

of the validity of the name used is often presented.

10. Etymology. The semantic roots or origin of the scientific name

are explained.

11. Description Usually just a subheading is given here.

12. Illustrations. All original illustrations in this report are listed

first, followed by references to selected illustrations of the species

available in the literature. The illustrations are not intended to portray

detailed anatomy (for example fin-ray counts) of the species, but rather

give an overall impression of the species' basic morphology.

13. Diagnosis. A brief diagnosis of each species is given.

14. Size. The maximum size, as well as the size range of specimens usually

encountered are given.

15. Morphology. This section provides a brief physical description of

the most notable anatomical features of each species that are not mentioned

in the diagnosis.






16. Counts. Fin ray and scale counts are usually given. In most cases

only the range is given, but sometimes detailed counts are presented as the

count followed by the number of fish counted; for example: AR: 22(9) means

a count of 22 anal-fin rays was obtained for nine specimens examined.

The following abbreviations are used: DR--Dorsal Rays; AR--Anal Rays;

PR--Pectoral Rays; VR--Ventral or Pelvic Rays; LS--Lateral Scales;

LLS--Lateral-Line Scales; PDS--Predorsal Scales; TS--Transverse Scales,

(this is sometimes divided into scales above and below the lateral line;

CPS--circumpeduncular scale count; GR--gill rakers (if not indicated as

the total count of both upper and lower limbs, the figure given is the

count of the rakers found on the lower limb of the outer gill arch).

17. Measurements. Common body measurements are presented. The following

abbreviations are used: SL--Standard Length; TL--Total Length; GBD--Great-

est Body Depth; PDL--Predorsal Length; HL--Head Length; EYE--Eye Diameter;

SNT--Snout Length; IO--Interorbital Distance.

18. Pigmentation. Life colors are usually given along with pigmentation

patterns that remain in preserved specimens.

19. Distribution and Natural History Comments that don't fit into other

sections are given here.

20. Range. The overall geographic range is given.

21. Apure Distribution. Map: fig. #. The number of the distribution map

for each species is given. In these maps, the collection localities are

plotted on a base map of the Apure drainage. Map scale sometimes required

that more than one locality be represented by a single dot. Brief comments

on the distribution within the Apure drainage are usually given. One should

bear in mind that dot maps of species distribution do not take into account

the fact that a few species in the Apure drainage are migratory, and thus







may be present in some areas during only part of the year. Also, non-

migratory species will spread out onto the floodplain in the wet season.

22. Habitat. A brief summary of the types of water bodies occupied by

the species is given.

23. Abundance. This is a somewhat subjective classification scheme, since

collection effort was not standardized to allow actual abundance data to be

calculated. I plotted the number of characid species versus the number of

collections, then arbitrarily drew separations between four categories as

follows:

RARE: If found in 1 to 5 collections.

UNCOMMON: If found in 6-25 collections.

COMMON: If found in 25-200 collections.

ABUNDANT: If found in more than 200 collections.

As indicated, these data are not quantitative, since fishing effort

varied considerably from site to site and not all specimens of each species

were preserved (especially true of the larger, more common species). Also,

the number of collections may include repeats from the same site. However,

the results are useful as a first approximation, and when used in conjunc-

tion with the distribution maps they usually reflect a fairly accurate

picture of relative abundance, or at least ease of capture.

24. Number of specimens examined. This is the total number of specimens

collected in the Apure drainage as part of this investigation. It does not

include specimens examined from other collections.

25. Food. Species are usually classified as CARNIVORE, HERBIVORE,

OMNIVORE or DETRITIVORE, and the major items that comprise each species

diet are listed if known. This section is based on my own field obser-

vations of live fishes and from stomach content analyses reported in




19

the literature, or done by myself, Dr. Craig Lilyestrom, Dr. Kirk

Winemiller or Mr. Leo Nico in our lab at UNELLEZ in Guanare.

26. Reproduction. Most of these data are from the literature, or

from data gathered by Dr. K. Winemiller for his doctoral research and

that was briefly summarized by Winemiller & Taphorn (1989). I use the

terminology introduced in that paper to classify the reproductive

strategy of each species:

"K-strategy" is characterized by high juvenile and adult survi-

vorship, low fecundity, a long life and generation time (the time from

birth or hatching to reproductive maturity), iteroparity (spawning

several times per season) and a relatively stable population density

throughout the year.

"rl-strategy" is characterized by low juvenile and adult survi-

vorship, low fecundity per individual but repeated spawning bouts

(iteroparity), a short life and generation time, and variable popula-

tion densities throughout the year.

"r2-strategy" is characterized by low juvenile but high adult

survivorship, a high fecundity, a long life, semelparity (spawning all

at once, once per season) and by great fluctuations in the population

density.

27. Migrations. There are no comprehensive studies of migration of the

fishes in the Apure Drainage, but we are currently investigating the migra-

tion and life cycle of the coporo, Prochilodus mariae (Barbarino D. &

Taphorn 1989). Local fishermen also provided information.

28. Importance. The commercial, sport, and ornamental uses as well as

the harmful or dangerous aspects of each species are discussed in this

section.




20

If no information is available on a given topic, that section is

omitted from the species account.

An abbreviated list of the material examined (the number of lots

or jars and the total number of specimens) is given for each species in

the Appendix. A more detailed list of specimens examined and locali-

ties is available from the author. All specimens are deposited in the

Collection of Fishes, Zoological Musuem, Museo de Ciencias Naturales de

Guanare, at UNELLEZ, in Guanare, Venezuela.

Each family is presented in a separate section, and appears in

alphabetical order. A brief diagnosis of the family, with comments on

the number of species and their general ecology is followed by a key to

the species of the Apure drainage and the individual species accounts.

In some keys additional information useful for identification, follows

the species name in brackets.













RESULTS


After many years of sampling (1977-1989), during which some 875

collections were made in the Apure drainage, over 14,000 lots of specimens

have been processed, cataloged and identified. I have identified 354

species of fishes, belonging to 41 families (Table 1). The so-called

ostariophysans strongly dominate the fauna, and comprise 305 (86%) of the

354 species. The ostariophysans have the anteriormost four or five verte-

brae fused and include, among South American groups, the orders Characi-

formes, Gymnotiformes and Siluriformes. The vast number of fishes col-

lected and data generated by the project have made it necessary to limit

this dissertation to a subset of the total fish fauna. The order Characi-

formes was selected because of its great diversity, and a somewhat larger

volume of literature that is available for identification of the species.

This order includes 13 (32%) of the 41 families and 138 (39%) of the

species (Table 1). Only the order Siluriformes, with about 137 species,

matches the characiform fishes in diversity in the Apure drainage. The

taxonomy of South American fishes is still relatively poorly known, and

has been likened to the situation in North America 100 years ago. As more

collecting is done, and taxonomic revision continues, the total number of

species per family will continue to change, but the relative numbers of

species per family will probably not vary as greatly, except perhaps in

the order Gymnotiformes, which will probably outstrip the others in terms

of new species described.







Composition of the Apure River Drainage Fishes.


Order

1 RAJIFORMES


2 CLUPEIFORMES


3 SALMONIFORMES*

4 CHARACIFORMES












5 SILURIFORMES


6 GYMNOTIFORMES





7 ATHERINIFORMES


8 SYNBRANCHIFORMES

9 PERCIFORMES


Family

1 Potamotrygonidae

2 Clupeidae
3 Engraulidae

4 Salmonidae

5 Anostomidae
6 Characidae
7 Characidiidae
8 Chilodontidae
9 Ctenoluciidae
10 Curimatidae
11 Cynodontidae
12 Erythrinidae
13 Gasteropelecidae
14 Hemiodontidae
15 Lebiasinidae
16 Parodontidae
17 Prochilodontidae

18 Ageneiosidae
19 Aspredinidae
20 Astroblepidae
21 Auchenipteridae
22 Callichthyidae
23 Cetopsidae
24 Doradidae
25 Hypophthalmidae
26 Loricariidae
27 Pimelodidae
28 Trichomycteridae

29 Apteronotidae
30 Electrophoridae
31 Gymnotidae
32 Hypopomidae
33 Rhamphichthyidae
34 Sternopygidae

35 Belonidae
36 Rivulidae
37 Poeciliidae

38 Synbranchidae

39 Cichlidae
40 Sciaenidae


10 PLEURONECTIFORMES 41 Soleidae


Species
Family
4


per
Order
4


% of Total
1.1


5 1.4


1 0.2


38.9












38.9


31 8.7


10 3.0


1 0.2

25 7.0


2 0.6


Table 1.







Characiformes

Characiform fishes are ostariophysans, the group of fishes that

dominates freshwater habitats in most of the world. Ostariophysans are

combined into a group because they all have the first few vertebra modi-

fied and linked to the surface of their bodies by a series of modified

bones to form what is presumed to be a listening device, or vibration

detector. Characiforms have gone farther with this device than most of

their cousins the catfishes, knifefishes, carps and suckers (the other

ostariophysans). They have developed a complete Weberian apparatus in

which the tripus is connected to the body of the third vertebra by means

of a vibrating lamina. This mechanism presumably transmits sounds or

vibrations picked up on the gas bladder to the inner ear (GUry 1977).

Better hearing could perhaps account in part for the dominance of the

ostariophysans in most freshwater ecosystems of the world (Fink & Fink

1981). Weitzman (1954, 1962) provided an extensive osteological defini-

tion of characiform fishes and Fink & Fink (1981) discussed their rela-

tionships with the other ostariophysans. The group is so diverse morpho-

logically that they tend to defy succinct definition. About all one can

say is that they all have scales and teeth on the jaws or lips! Actually,

not even that is true of all characiforms since a few species from the

high Andes in southern South America have lost their scales completely and

the Curimatidae are toothless (at least as adults; juveniles have teeth in

the early developmental stages). Many are small schooling species that

have a chemoreceptory sensory system that, upon detection of substances

released by school members under attack or stress, will trigger a "fright

response," causing the school to adapt defensive maneuvers.







No consensus has yet been reached by workers on characiform upper-

level taxonomy. For many years almost all characoids were lumped in the

family Characidae (Eigenmann 1917, Weitzman 1962). The group was subdivided

by Greenwood et al. (1966), which (for Characiformes) is based largely on

the findings of Dr. Stanley Weitzman. Since then, the group, and the divid-

ing lines between the families and subfamilies, have been topics of constant

investigation, and debate. Gdry (1977) proposed several modifications. In

Table 2 the two schemes are contrasted for South American groups.


Table 2. Comparison

Greenwood et al.

(12 families)

Characidae

Characidae

Characidae

Characidae

Erythrinidae

Ctenoluciidae

Cynodontidae

Lebiasinidae

Parodontidae

Gasteropelecidae

Prochilodontidae

Curimatidae

Anostomidae

Hemiodontidae

Chilodontidae


of family level taxonomy of Characiformes.

Gdry Used Here

(11 families) (13 families)

Characidae Characidae

Characidiidae Characidiidae

Serrasalmidae Characidae

Crenuchidae Characidae

Erythrinidae Erythrinidae

Ctenoluciidae Ctenoluciidae

Characidae Cynodontidae

Lebiasinidae Lebiasinidae

Hemiodidae Parodontidae

Gasteropelecidae Gasteropelecidae

Curimatidae Prochilodontidae

Curimatidae Curimatidae

Anostomidae Anostomidae

Hemiodidae Hemiodontidae

Curimatidae Chilodontidae







Thus, the scheme of Greenwood et al. (1966) would leave South America

with 12 families of Characiformes to Gdry's 11. In more recent works, some

of these families have been promoted to familial or demoted to subfamilial

rank. Weitzman & Gdry (1981) defined the Characidiinae. Nelson (1984)

listed Cynodontinae (which he earlier [1976] recognized as a family), as a

subfamily of Characidae; Vari (1989) treated it as a tribe of the Characi-

dae. Other groups have had the familial limitations better clarified and

justified, as for example Vari's (1983, 1989) treatment of Curimatidae,

Prochilodontidae, Anostomidae and Chilodontidae. Nelson (1984) included

only eight families: Hemiodontidae, Curimatidae, Anostomidae, Erythrini-

dae, Lebiasinidae, Gasteropelecidae, Ctenoluciidae and Characidae, thus

demoting again the Prochilodontidae, Cynodontidae, Parodontidae and

Chilodontidae. Changes will continue. Except for the recognition of the

family Characidiidae, I have followed the classification of Greenwood et

al., for a total of 13 families. The correct spellings of Hemiodontidae

and Chilodontidae (vs Hemiodidae and Chilodidae) are also as yet unre-

solved. The type genera are Hemiodus and Chilodus.

I have adopted this system as a matter of practical convenience. This

should not be interpreted as support for any particular position in the

controversy of higher level relationships of characiforms. For convenience,

and since taxonomic affinities are as yet unresolved at various levels, the

families, genera and species are discussed in alphabetic rather than taxo-

nomic order.

The characiform fishes exhibit an incredible diversity of form

and occupy a wide array of ecological niches. Eigenmann (1917), Weitz-

man (1962) and Fink & Fink (1981) all marveled at the incredible explo-

sive radiation of this group in South America, which surpasses that of







the marsupials in Australia. Although many families contain widely

diverging morphotypes, I will try to briefly characterize each.

The Characidae comprises the most morphologically diverse group and

also includes the largest number of species. Although most species are

small minnow-like tetras ("sardinas" in Venezuela), this family includes

the piranhas, giant pacus, pike characins, strange tusked scale-eaters,

tiny killyfish-like inhabitants of blackwater streams, neon tetras, silver

dollars and palometas, and South American characid versions of trout,

salmon, clupeids and anchovies. The Erythrinidae is a small family of

large to medium fishes that are cylindrical, bowfin-like predators, typi-

cal of shallow lentic habitats. The Ctenoluciidae is represented by only

a few species of elongate gar-like surface predators. Cynodontids are

large, riverine predators with keeled, compressed bodies, large eyes and

expanded pectoral fins adapted to life near the surface in strong cur-

rents. Their extremely large canines give these piscivores a particularly

fierce appearance. Lebiasinids include two divergent groups: the "Creole

carps," Lebiasina, are medium-sized predators that inhabit mountain

streams, whereas Pvrrhulina and relatives are small, colorful fishes

typical of vegetation-choked lowland ponds. The Characidiidae and Paro-

dontidae include ecological equivalents of North American darters, adapted

to life in the riffles or mountain streams. The hatchetfishes of the

family Gasteropelecidae are bizarre, insectivorous, surface-dwelling

fishes with incredibly expanded coracoid bones (Weitzman 1954, 1960), and

very large pectoral-fins that, when rapidly vibrated, give these fish the

ability to "fly." Prochilodontids and curimatids are medium to large

sized fishes reminiscent of carps and suckers. They feed on mud, algae

and detritus, and comprise a large percentage of the biomass in most







tropical waters. They frequently support productive local commercial

fisheries. Many species make spectacular annual migrations to feeding or

breeding grounds. Anostomids and hemiodontids include mostly herbivores

and planktivores, with fusiform bodies and conical heads adapted to swim-

ming in fast currents. The chilodontids are small to medium-sized head-

standers similar to the curimatids, but with teeth.

The differing, still varying taxonomic classifications of characi-

forms exist primarily because current taxonomic groups for the most part

reflect different authors' attempts to group morphologically similar taxa

together. Differing opinions as to exactly which characters are "similar"

and which are not leads to different conclusions. Although Eigenmann

(1917) expressed much concern about the monophyly of the numerous upper

level taxa he described, the methodologies available at that time were

inadequate. He remarked that the tremendous "radial adaptation" of char-

aciform fishes was at once a paradise for evolutionary biologist and a

nightmare for taxonomists seeking a useful system of names. Durbin Ellis

(1918) came to the conclusion that the genera Hyphessobrvcon and Hemiqram-

mus were conveniences, and not natural entities (ie. monophyletic units).

Recent efforts, such as those as Vari (1989), to base taxonomy on shared

derived characters and that reflect the phylogenetic history of the fishes

would seem to be the only approach that might eventually lead to agreement

among taxonomists instead of continuing controversy. The characiform

families, and especially the Characidae, contain such divergent groups of

fishes that it is difficult to write a key that is both easy to use and

reasonably precise. The following key should work for most individuals of

most species currently known from the Apure drainage. Readers are also

referred to Taphorn & Lilyestrom (1984) for keys to all the freshwater

families in Venezuela.







Key to the Characiform families present in the Apure River drainage.

la. Lips and jaws completely lacking teeth of any kind...

CURIMATIDAE

lb. Teeth present on jaws or lips (teeth on lips may be small and

difficult to see without magnification)... ...2

2a. (Ib) Teeth present only on lips, none on jaws ...3

2b. Lips without teeth; upper or both jaws with teeth... ...4

3a. (2a) Gills normal; mouth large and evertible to form a round,

sucking disk; teeth present on edges of lips, teeth small, comb-

like and numerous (many more than 20 on either side), arranged in

single row at sides of lips, and in two rows near center midlinee)

of lips; predorsal spine present, embedded in flesh just anterior

to dorsal fin; large species that exceed 200 mm SL in the first

year of growth... PROCHILODONTIDAE

3b. Gills with fourth arch dilated (expanded), its surface with folds

and wrinkles that mesh with fifth arch; mouth small, not complete-

ly evertible; teeth larger, not very numerous (less than 20 per

jaw), in single row; no predorsal process present; small species
that seldom exceed 200 mm SL CHILODONTIDAE

4a. (2b) Lower jaw without teeth, or without teeth at center (near

symphysis)... ...5

4b. Lower jaw with complete series of teeth... ...6

5a. (4a) Teeth of upper jaw arranged in semicircle; pectoral fins

not greatly expanded, usually not as long as head; cranium without

fontanels; lower jaw never with teeth at sides, usually completely

lacking teeth... HEMIODONTIDAE







5b. Teeth of upper jaw arranged in straight line; pectoral fins

greatly expanded, usually longer than head; cranium with large

fontanels; lower jaw usually with a few teeth at sides...

PARODONTIDAE

6a. (4b) Lower jaw trapdoor-like, with extremely large canine teeth,

(the length almost equal to or surpassing eye diameter) that fit

into special "sockets" present in upper jaw and cranium when mouth

is shut... CYNODONTIDAE

6b. Lower jaw without extremely large canines, sometimes with small

canines that measure less than one third of eye diameter, but

usually with multicuspid teeth... ...7

7a. (6b) Adipose fin present... ...8

7b. Adipose fin absent... ...13

8a. (7a) Scales ctenoid; jaws extremely elongate and anal fin with

fewer than fifteen rays... CTENOLUCIIDAE

8b. Scales cycloid; jaws usually not extremely elongate, but if so,

anal fin with more than 15 rays... ...9

9a. (8b) Branquial membranes fused to isthmus for nearly their entire

length; mouth with 6-8 large rabbit-like, sometimes unusually

shaped incisors... ANOSTOMIDAE

9b. Branquial membranes free from isthmus; teeth variable, but not

usually as above... ...10

10a. (9b) Chest bones (coracoids) greatly expanded to form a deep

narrow keel; dorsal-fin origin behind that of anal fin; lateral

line short, usually directed down towards anal fin origin; pec-

toral fins greatly enlarged for flight...

GASTEROPELECIDAE







10b. Chest usually not greatly expanded, but if so, then dorsal-fin

origin is at a level with, or in front of that of anal fin;

lateral line, if present not usually directed down towards

anal-fin origin; pectoral fins variable... ...11

Ila (lOb) Lower jaw with two rows of tricuspid teeth (the second row

of small teeth sometimes difficult to see); upper jaw with only

one row of tricuspid teeth and without conical or canine teeth...

LEBIASINIDAE (Lebiasina)

lib. Teeth not as described above... ...12

12a. Pectoral and pelvic fins enlarged, inserted very low on body and

directed under chest (to support fish resting on substrate);

pectoral fin with the first three or four rays thickened and

unbranched; premaxillary teeth always in a single row; chest and

abdomen flattened (adapted to benthic life as in North American

darters); head conical; mouth small usually subterminal to ventral

(but terminal in a few species)...CHARACIDIIDAE (in part)

12b. (lib) Pectoral and pelvic fins not particularly expanded, usually

inserted near or just below midbody, (pectorals not used to

support body when resting on substrate); first three or four rays

not thickened, usually only first one or two unbranched; premaxil-

lary teeth in one to three rows, but usually two; chest and abdo-

men rounded; head not usually conical; mouth variable...

CHARACIDAE (in part)

13a. (7b) Large (up to 1000 mm SL) predators with wide terminal mouths

armed with canine teeth on premaxilla and dentary, and sometimes

on maxilla; scales relatively large (some nearly size of eye)...

ERYTHRINIDAE







13b. Small (less than 40 mm SL) fishes with small, superior or

terminal mouths that lack large canine teeth (teeth usually small

to moderate in size, conical or multicuspid); scales relatively

small... ...14

14a. (13b) Dorsal fin origin posterior to that of anal fin; males

with long filamentous extension on opercle...

CHARACIDAE (Corvnopoma)

14b. Dorsal fin origin anterior to that of anal fin; males lacking

opercular filament... ...15

15a. (14b) Mouth terminal or subterminal; dorsal fin with fifteen or

more rays; sides with alternating light and dark vertical bars...

CHARACIDIIDAE (Elachocharax)

15b. Mouth superior; dorsal fin with fewer than fifteen rays; sides

without vertical bars... LEBIASINIDAE (pyrrhulinines)

Anostomidae

The anostomids characteristically are fusiform, somewhat elongated

fishes with conical heads, distinguished from other families by their small,

nonprotractile mouths that usually have eight large incisors arranged in a

single row in either jaw. The anterior nostril is usually tubular, and the

gill membranes are often united to the isthmus (Gdry 1977). According to

Winterbottom (1980), this family together with the Chilodontidae, shares

posteriorly replaced, cuspid pharyngeal teeth (not to be confused with the

pharyngeal teeth of Cyprinidae). Roberts (1969), Gery (1977), Winterbottom

(1980) and Vari (1983) have presented very different concepts of subfamilial

divisions within the Anostomidae. The family comprises about ten genera,

many of which are monotypic. All told, there are at least 100 described





32

species (Gdry 1977), of which the largest by far is Leporinus, with some 70

nominal species (Vari 1983).

Most of the remaining taxonomic problems in this family involve the

genus Leporinus, since Winterbottom's (1980) work has cleared up the

situation in the subfamily Anostominae (comprising the genera Anostomus,

Pseudanos, Gnathodolus, Sartor, and Svnaptolaemus). Neither he nor Vari

(1983) defined for the remaining genera, Abramites Abramoides, Laemolvta,

Leporellus, Leporinus, Rhytiodus and Schizodon, although Winterbottom

(1980) rejected Gery's (1977) subdivision into two subfamilies, Leporelli-

nae for Leporellus, and Anostominae for all the rest.

The majority of anostomids are medium to large-sized omnivores or

vegetarians that use their incisor-like teeth to clip aquatic vegeta-

tion, or as forceps to remove prey from crevices. A few anostomids

have developed feeding specializations, these species are relatively

smaller in size, are more elongated and have minute upturned mouths,

often with bizarre dentition.

I have been able to identify twelve species from the Apure drainage,

half of them members of the genus Leporinus. Most of the species'

identifications are still tentative. For those that don't seem to agree

with any described species, I have used code names.

Very little natural history information is available on anosto-

mids in the Apure drainage. Most species are probably omnivorous,

though some like Schizodon isoqnathus tend to be more herbivorous. All

are probably egg scatterers, with no parental care. It is expected

that most of the medium to large species spawn annually at the onset of

the rains. The smaller ones probably spawn repeatedly during the wet

season.







Key to the Apure Drainage Species of Anostomidae

This key is based in part on Winterbottom (1980) and Gery (1977),

and will work only for specimens with adult pigmentation patterns.

Since there is considerable size range variation in the family, defining

"juvenile" size for all anostomids is not possible. Juvenile Anostomus

for example, probably obtain adult coloration at a size of 20-30 mm SL,

whereas juvenile Leporinus may not change to the adult pattern until

they reach 100-120 mm. Juveniles all tend to look alike, and are

marked with numerous vertical bars and rounded midlateral spots, that

are gradually lost as the adult pattern is expressed.

la. Caudal fin with oblique black bars; caudal fin densely scaled

for most of its length; anterior nostril not tubular, proximate

in position to the posterior nostril...

Leporellus vittatus (fig. 7)

lb. Caudal fin uniformly pigmented, without bars; caudal fin not

scaled; anterior nostril usually tubular and well separated

from posterior nostril... ...2

2a. (Ib) Mouth superior (i.e. upturned) in adults... ...3

(Note: small juvenile Schizodon isognathus have superior mouths

to a size of some 4 cm SL.)

2b. Mouth terminal or inferior in adults... ...5

3a. (2a) Pigmentation pattern comprising a large rounded black spot

surrounded by lighter area located on center of each side, a

smaller spot behind opercle and at base of caudal fin, dorsum

crossed by 8-11 irregular narrow black bars, sides with horizontal

rows of small spots... Pseudanos irinae (fig. 23)







3b. Pigmentation pattern not comprising a large rounded spot in center

on the side, but rather of several wide black stripes or one wide

black lateral stripe with several additional horizontal rows of

small dots ...4

4a. (3b) Pigmentation pattern consisting of 10-11 horizontal rows of

small dots on sides and a wide lateral stripe from opercle to tail

(fig. 21) Pseudanos gracilis (fig. 21)

Winterbottom (1980) noted that this species exhibits two color

phases. One is described here, whereas the other (not yet found

in the Apure drainage) consists of 3-4 large rounded spots on mid

sides with horizontal rows of small whitish spots. This is simi-

lar to the pattern of P. irinae; however, P. gracilis never has

horizontal rows of black dots nor dark thin bars crossing the

dorsum.)

4b. Pigmentation pattern consisting of wide black horizontal stripes

separated by narrower white stripes, the widest black stripe

centered on the side and extending from opercle to base of tail...

Anostomus ternetzi (fig. 5)

5a. (2b) Teeth incisiform, with 2-5 cusps, some multicuspid teeth

always present; anal fin usually with only seven branched rays...

Schizodon isognathus (fig. 25)

5b. Teeth incisiform, without cusps or at most bifid, never

multicuspid; anal fin with eight or more branched rays... ...6

6a. (5b) Anal fin usually with ten or more branched rays; body high

and compressed, GBD 37-48% SL; a postventral keel present;

headstanders that usually swim in a vertical position...

Abramites hypselonotus (fig. 3)







6b. Anal fin usually with nine branched rays or fewer; body terete, not

very compressed, GBD usually less than 30% SL; no postventral keel;

swimming position horizontal... (genus Leporinus, 6 spp.)... ...7

7a. (6b) Body with about seven, wide, black vertical bars, the second

usually forming a "Y"; base color yellowish...

Leporinus yophorus (fig. 19)

7b. Body without vertical bars, either plain, with horizontal stripes,

or with large spots; base color variable... ...8

8a. (7b) Body with 4-5 wide black horizontal stripes, separated by

narrower white or yellow stripes...

Leporinus cf striatus (fig. 17)

8b. Body without horizontal stripes, either plain with a small caudal

spot or with large rounded spots on sides... ...9

9a. (8b) Adults with plain brown body and a small black spot at base

of caudal fin... Leporinus sp. (fig. 13)

[lateral-line scales 42-44; predorsal scales 12-13; transverse

scales 13-14]

9b. Adults with large black spots on sides of body... ...10

10a. (9b) Adults with five or more large black spots on sides of body,

some along lateral midline, others both above and below these;

mouth inferior... Leporinus cf maculatus (fig. 9)

1Ob. Adults with only two to four large rounded spots, all along lateral

midline; mouth terminal or subterminal... ...11

11a. (lOb) Adults usually with only two large rounded spots along

lateral midline and no faint row of smaller spots on upper sides,

although faint vertical saddles cross dorsum; head and body deeper

and more robust; scales seldom spotted (one spot per scale); body







often decidedly higher in front of dorsal fin; eye smaller, in

specimens over 100 mm SL its horizontal diameter less than half of

interorbital width... Leporinus friderici (fig. 11)

[lateral-line scales 38-40; predorsal scales 9-12; transverse

scales 11-12]

11b. Adults usually with three or four large rounded spots along

lateral midline plus a faint row of smaller spots on upper sides;

scales often a spot (one per scale) arranged so as to form hori-

zontal rows of dots; head and body more slender, not decidedly

higher in front of dorsal fin; eye larger, in specimens over 100

mm SL its horizontal diameter is more than half interorbital

width... Leporinus sp. "aguaro" (fig. 15)

[lateral-line scales about 39; predorsal scales 10-12; transverse

scales 11-12]

Species Accounts

Abramites hypselonotus (Gunther) 1868

High-Backed Headstander Cabezibajo

Fig. 3. Map: fig. 4. Couplet 6a.

Generic synonymy

Abramites Fowler 1906:331 (type species: Leporinus hypselonotus GUnther

1868, by original designation); Vari & Williams 1987:89 (revision of

genus).

Specific synonymy

Leporinus hypselonotus GUnther 1868a:480 (type locality: Upper Amazon,

Xeberos, Peru); Steindachner 1882:12 (Ciudad Bolivar, Venez.);

Eigenmann 1909:323, 344 (Orinoco); Schultz 1944b:268 (Venez.).








































Figure 3.


Abramites hypselonotus.







Leporinus solarii Holmberg 1887:222 (type locality: Argentina, Rio

Parani).

Leporinus eaues Boulenger 1896:34 misidentification of L. hypselonotus

from Brazil).

Abramites hypselonotus Fowler 1906:331 (indicated as type species of

Abramites; 1950:249 (synonymy); Mago L. 1970:75 (Venez.); G6ry

1977:175 (key); Vari & Williams 1987:92 (synonymy, key, revision of

genus).

Abramites microcephalus Norman 1926:92 (type locality: near mouth of

Amazon River).

Abramites ternetzi Norman 1926:93 (type locality: Brazil, Matto (= Mato)

Grosso, Sao Luis and Descalvados.

Leporinus salarii Borodin 1929:288, (as a possible synonym of L.

hypselonotus, specific name misspelled).

Leporinus nigripinnis Meinken 1935:193 (type locality: Argentina,

Corrientes).

Abramites eaues FernAndez Y. 1950:116 misidentificationn, Rio

Salinas, Venez.); Mago L. 1970:75 (Venez.).

Abramites solarii Ringuelet, Aramburu and Alonso de Aramburu 1967:213.

Abramites hypselonotus ternetzi Gery 1977:175 (Rio Paraguay Basin).

Abramites hypselonotus hypselonotus Gdry 1977:75 (Amazon and Orinoco

basins).

Types. (designated by Vari & Williams 1987). Lectotype: BMNH

1867.6.13:40. Paralectotypes: BMNH 1867.6.13:41-42.

Comments. The genus Abramites is comprised of only two species. A.

hvyselonotus (GQnther 1868), the only species in Venezuela, has 10-12

branched anal-fin rays and eight, irregularly-shaped bars on the sides







between the nape and the caudal peduncle. A. eaues (Steindachner) 1878,

has 13-14 branched anal-fin rays and only five bars on the body, the

anteriormost situated under the dorsal fin (Vari & Williams 1987).

Etymology. ABRAM = from Abramis, a genus of similarly shaped fish,

ITES = like; HYPSELO = raised or high, NOTUS = back.

Description

Illustrations. Fig. 3; Axelrod et al. 1971:F-2.10; Vari & Williams

1987:figs. 2-5.

Diagnosis. The unusual body shape and pigmentation pattern (fig. 3)

are sufficient to identify this species. It is further distinguished by

a terminal mouth, a deep, compressed body (GBD 37-48% of SL), and the

presence of a postventral keel.

Size. It grows to about 130 mm SL.

Morphology. See Vari & Williams (1987). The conical head abruptly

expands vertically behind the eye and the dorsal profile rises steeply

from the nape to the dorsal-fin origin to give this species its charac-

teristic high-backed appearance. The body is more compressed than in

most anostomids.

Counts. DR 10-13 (up to three unbranched); AR 12-15 (up to three

unbranched); PR 13-15; VR 8-9, 37-38 vertebrae.

Pigmentation. The body is tan or silvery with eight irregular

vertical bars, the darkest and widest bar extends from the pelvic fins

across body and onto the anterior dorsal fin. There is also a black

stripe from the tip of the lower jaw through the eye, which continues

obliquely upward toward the dorsum. The adipose fin is yellow and

black, the caudal and pectoral fins are clear, and the anal fin is

black.

















































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Distribution and Natural History

Range. A. hypselonotus occurs in the Amazon, Orinoco, Paraguay and

lower Parand basins (Vari & Williams 1987).

Apure distribution. Map: fig. 4. Although few specimens were

collected, the localities range across the entire Apure drainage.

Habitat. This species was most frequently collected in habitats

with aquatic vegetation in quiet sidewaters of large whitewater rivers

in the low llanos. It swims in a head down position, often aligning

itself with the stems of plants.

Abundance. UNCOMMON.

Number of specimens examined. 28 from 14 collections.

Food. OMNIVORE. Its diet includes aquatic invertebrates, as well

as vegetable matter.

Reproduction. Probable strategy: rl.

Importance. This is a highly valued ornamental.

Anostomus ternetzi Fernandez Y. 1949

Red-Mouthed Headstander Cabezibajo Bocaroja

Fig. 5. Map fig. 6. Couplet 4b.

Generic synonymy

Anostomus Gronow 1763:122 (description and figure, no species mentioned,

work not accepted by ICZN).

Anostomus Scopoli 1777:451 (no species mentioned, but since this work

was based on Gronow the type species is that illustrated by Gronow

1756, which is Salmo anostomus Linnaeus).

Anostoma Rafinesque 1815 (alternative spelling).

Mormyrhynchus Swainson 1839:186, 291 (type species: Mormyrhynchus

gronovii Swainson = Salmo anostomus Linnaeus, by monotypy).







































































Figure 5. Anostomus ternetzi.


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Histriodromus Gistel 1848:8 (type species: Salmo anostomus Linnaeus,

by monotypy).

Pithecocharax Fowler 1906:319 (type species: Salmo anostomus Linnaeus,

by original designation, this name was proposed as a substitute for

Anostomus Walbaum 1792).

Specific synonymy

Anostomus ternetzi Fernandez Y. 1949:293 (type locality: Orinoco River

System); Knoppel 1972:231 (diet).

Anostomus anostomus (non Linnaeus); Myers 1950:184 (part).

Types. Holotype: Museo de Ciencias Naturales de Caracas 46001.

Paratypes: MCNC 46002; CAS 20093(1), 20094(1), 20095(4), 20096(1).

Etymology. ANO = upturned, STOMUS = mouth; TERNETZI = for Carl

Ternetz.

Description

Illustrations. Fig. 5; Axelrod et al. 1971:F-40.11 40.12; Gdry

1977:188; Winterbottom 1980:81 fig. 17.

Diagnosis. A. ternetzi is separable from Leporinus by the superior

mouth. The color pattern is distinctive for the species. The white

stripe that extends from between nostrils along the dorsal midline to

the dorsal-fin origin in this species, is dark in A. anostomus, a spe-

cies that occurs in other parts of Venezuela.

Size. It grows to about 100 mm SL.

Counts. This species has three branchiostegal rays instead of the

four usually found in other Anostomus species. DR 12-13; AR iii7-iii8;

PR i13-i15; VR i8; LS 39-42; PDS 11-14; CPS 16.

Pigmentation. The basic pigmentation pattern consists of three wide

black horizontal stripes separated by narrower white zigzag stripes.







The widest black stripe is situated at midside and extends from the

opercle to the base of the tail. Another starts at the pectoral-fin

insertion and extends back to the lower part of the caudal peduncle.

There is a narrow black stripe situated on either side of the narrow

white stripe that runs along the ventral midline, and another wide black

stripe on either side of the white stripe that extends from between the

nostrils to the dorsal fin origin. In many individuals the black (or

dark brown) pigment uniformly covers most of the dorsum, although it is

sometimes divided into several rows of dots by lighter areas. Fins that

are normally reddish in life, would appear clear in preserved specimens.

Distribution and Natural History

Range. This species occurs in the Amazon and Orinoco basins, and in

Guyana.

Apure distribution. Map: fig. 6. It is known only from northern

Apure state, and the Aguaro-Guariquito system.

Habitat. It is found in black or clearwater streams with abundant

aquatic vegetation, in the sandy-soiled areas of northern Apure state

and western Guirico.

Abundance. UNCOMMON.

Number of specimens examined. 69 from 19 collections.

Food. OMNIVORE. This fish probably feeds on vegetable material as

well as aquatic insects. It swims head down, and presumably feeds in

this position amidst aquatic vegetation.

Reproduction. Probable strategy: rl. Winterbottom (1980) noticed

some sexual dimorphism, in that males have a greater caudal peduncle

depth. Sterba (1972) stated that the similar A. anostomus is territori-

al. Axelrod et al. (1971) stated that in A. anostomus the pair quivered

























































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side by side, and expelled a large number of eggs all over the glass,

plants and the bottom of the aquarium. Fry were slender, about 15 mm

long, and swam in a normal horizontal position.

Importance. Ornamental.

Leporellus vittatus (Valenciennes) 1849

Stripe Tailed Leporellus Mije de Cola Rayada

Fig. 7. Map fig. 8. Couplet la.

Generic synonymy

Leporellus Lutken 1874b:129 (type species: Leporinus pictus Kner, by

monotypy).

Leporinodus Eigenmann 1922:116 (type species: Leporinodus retropinnus

Eigenmann 1922, by original designation, (described in footnote).

Specific synonymy

Leporinus vittatus Valenciennes in Cuvier & Valenciennes 1849:59 (type

locality: Rio Amazonas).

Leporellus vittatus Lutken 1874b:129; Fowler 1950:253 (synonymy);

Gery 1977:151 (diagnosis).

Leporinodus vittatus Eigenmann 1922:117.

Leporinus pictus Kner 1859:172, pl. 8, fig. 19 (type locality: Orissanga,

Est. de SAo Paulo).

Salmo cagoara Natterer in Kner 1859:172 (name in synonymy).

Leporinus maculifrons Reinhardt in LQtken 1874b:201 (name in synonymy).

Leporellus timbore Eigenmann 1922:117 (type locality: Rio das Velhas).

Comments. Gdry (1977) listed seven forms in this genus, of which

he listed (without explanation) L. maculifrons, L. timbore, and L.

cartledqei as synonyms of L. pictus. None are well known, and a revi-

sion of the genus is necessary before any further comments can be made















































Figure 7. Leporellus vittatus.


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about synonymies. I have used the synonymy of Fowler (1950) here, but

additional study may change these.

Etymology. LEPORELLUS = like a rabbit, referring to the teeth;

VITTATUS = spotted.

Description

Illustrations. Fig. 7; Gdry 1977:153.

Diagnosis. This is the only anostomid with a scaled, obliquely

barred caudal fin. The pigmentation pattern of the body (fig. 7) is

also distinctive.

Size. It grows to at least 200 mm SL.

Morphology. It is elongate with a large head and eye. The mouth is

subterminal with the lower lip reverted to form a wide fold. The gill

membranes are joined to the isthmus. The nostrils are proximate, and

not tubular (unusual in this family). The caudal fin is completely

scaled.

Counts. DR 13; AR 10-11; PR 16-17; VR i8; LS 42; PDS 12-13; CPS

16-17; TS 11-12.

Pigmentation. The base color of the body is yellow-brown in life.

The white tail has a black stripe through the central rays, and two

black oblique bars in each lobe. The top of the head, dorsum, and upper

sides are heavily spotted, but the belly is white. The spots on the

sides (one per scale) are arranged in nine or ten horizontal rows. The

sides have two white stripes devoid of spots that continue posteriorly

onto the tail. The dorsal fin is white with a large black distal blotch

on its first rays, and a row of black spots nearer base. The pectoral

and pelvic fins are whitish to clear. The anal fin is white with a

central black blotch.



























































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Distribution and Natural History

Range. The range is not well documented but it probably occurs in

most of the Andes in Venezuela and Colombia. There are reports of this

species from the Guyana Shield in Venezuela, but I have not seen speci-

mens. The form there may represent a different species.

Arure distribution. Map: fig. 8. It has been found only in the

western portion of the Apure drainage, usually in the Andean foothills.

Habitat. It lives in clear, clean, fast-flowing mountain streams.

Juveniles have been taken at Bruzual in the quiet edges of the Apure

River from aquatic vegetation. They perhaps were washed out of the

mountains, or the reproductive cycle might include a downstream phase

with later upstream migration.

Abundance. UNCOMMON.

Number of specimens examined. 49 from 16 collections.

Food. OMNIVORE. It feeds on both insects and vegetable matter. I

observed several individuals biting at rocks in the clear waters of the

upper Rio Caparo, presumably selecting items from the epibenthos.

Reproduction. Probable strategy: r2.

Importance. It has great potential as an ornamental.

Leporinus cf maculatus MUller & Troschel 1844

Spotted Leporinus Mije Pintado or Cabeza de Manteco

Fig. 9. Map: fig. 10. Couplet 10a.

Generic synonymy

Leporinus Spix in Agassiz 1829:65 (type species: Leporinus fasciata

Agassiz, by monotypy).

Hypomasticus Borodin 1929:287 (type species: Leporinus mormvrops

Steindachner 1875, designated by Fowler 1950:228).






































Figure 10. Leporinus cf maculatus.







Specific synonymy

Leporinus maculatus Muller & Troschel 1844:86 (type locality: Guyana);

1845:11 (rivers of Guyana); Eigenmann 1912a:305 (redescription);

Fowler 1950:237 (synonymy); Gdry 1977:166 (key, discussion of many-

spotted Leporinus).

Leporinus meqalepis Gunther 1863:443 (type locality: Essequibo River,

Guyana).

Leporinus marcqravii Lutken 1874b:130 (type locality: Rio Velhas and

tributaries).

Comments. The counts for Apure drainage specimens are consistently

higher than those given by Eigenmann (1912) for specimens from Guyana,

either because of a more inclusive counting method, or because different

species are involved. Gery (1977) discussed the complex taxonomic

problems of the many-spotted Leporinus species. He stated that in

current usage both aquarists and ichthyologists have confused L. pelle-

grini with L. maculatus, and that the name L. maculatus has been applied

to at least two very different species. I follow him in using L. macu-

latus for the species with a ventral mouth. He placed this species in

the subgenus Hypomasticus.

Etymology. LEPO = rabbit, RINUS = snout; MACULATUS = spotted.

Description

Illustrations. Fig. 9; Eigenmann 1912:pl. 43, fig. 2; Axelrod et

al. 1971:F377.00; Gery 1977:166; RomAn 1985:141.

Diagnosis. The pigmentation pattern is distinctive (fig. 9).

This is the only Leporinus with five or more large spots on the sides.

The ventral mouth is also unique.

Size. It grows to at least 200 mm SL.







Morphology. The head length is about equal to the greatest body
depth. The interorbital width is less than the snout length.

Counts. DR iilO; AR ii9; PR i14-i16; VR i8; CPS 16; LS 38-40; TS
11-12 5-6 above, and 5 below lateral line.

Measurements. HL = 22% SL; GBD = 27-33% SL (Eigenmann 1912 for

Guyanan specimens). In Apure specimens, GBD and HL = 24-26% SL; EYE =

50-66% HL, 10 = 43-55% HL, and SNT = 20-24% HL.

Pigmentation. The body is marked with several large ovate black

spots along the lateral midline, and on the upper and lower sides This

is a pattern similar to that of most juvenile anostomids, but in this

case it has been retained in the adult.

Distribution and Natural History

Range. It occurs in the Guianas and in the Orinoco Basin.

Apure distribution. Map: fig. 10. It is known only from the Agua-
ro-Guariquito system in the easternmost portion of the Apure drainage.

Habitat. It occurs in clear and blackwater creeks of the upper
Aguaro River system, in sandy bottomed savanna streams with abundant

aquatic vegetation.

Abundance. RARE.

Number of specimens examined. 6 from 4 collections.

Food. HERBIVORE. It is probably mainly herbivorous, though occa-

sionally aquatic insects may be included in the diet.

Reproduction. Probable strategy: r2. It is probably an egg scatterer

that synchronizes its annual reproductive effort with the beginning of the

rainy season. Axelrod et al. (1971) reported that this species spawned in

aquaria, scattering eggs all over, and then eating them if not removed.

Importance. Ornamental.





















































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Leporinus friderici (Bloch) 1794

Leporinus Mije

Fig. 11. Map: fig. 12. Couplet 11a.

Specific synonymy

Salmo friderici Bloch 1794:94, pi. 378 (type locality: Surinam).

Leporinus friderici Muller & Troschel 1844:87; 1845:11; Eigenmann &

Allen 1942:305 (Venezuela); Fowler:1950:233 (synonymy).

Leporinus fridericii Kner 1859:170 (misspelling).

Leporinus frederici Valenciennes in Cuvier & Valenciennes 1849:25

(misspelling).

Leporinus fradericii Goeldi 1898:482 (misspelling).

Curimatus frederici Perugia 1891:641.

Curimatus acutidens Valenciennes 1847:9, pl. 8, figs 1-la (type locali-

ty: South America)

Leporinus megalepis (non Gunther 1863) Gunther 1864:307 (part).

Leporinus leschenaultii Valenciennes in Cuvier & Valenciennes 1849:30,

pl. 635 (type locality: Mana); Peters 1877:472 (Calabozo, Venezuela);

Eigenmann & Eigenmann 1891:51 (Calabozo, Venezuela); Pellegrin

1899:157 (Apure River, Venezuela).

Comments. Identification of species in the L. friderici group is

difficult, see Gdry (1977) and Garavello (1988) for a brief outline of

the problem. Apure specimens resemble L. subniqer Fowler 1943 from

Florencia, Colombia in the Andean piedmont, and both of these seem

quite similar to L. bahiensis Steindachner 1875 from Brazil.

Etymology. FRIDERICI = probably named for the German monarch

Frederick the Great (1712-1786).





































Figure 11. Leporinus friderici.







Description

Illustrations. Fig. 11; Eigenmann 1912:pl. 63 fig. 4; Axelrod et

al. 1971:F-376.00; Gdry 1977:168.

Diagnosis. The pigmentation pattern is distinctive (fig. 11). Adults

usually have only two large spots on the sides. This species is very simi-

lar to Leporinus sp. "aguaro," but that form usually has three large spots

on the sides, and has a relatively larger eye (the eye diameter is more than

half of the interorbital width vs less than half in L. friderici. Faded

specimens of L. friderici (without spots) are similar to "Leporinus sp.,"

but the latter has much wider lips, and more lateral-line scales (42-44 vs

38-40). L. boehlkei Garavello 1988 has fewer lateral-line scales (35-36),

and fewer circumpeduncular scales (12 vs 16).

Size. It reaches at least 350 mm SL.

Morphology. This is a very robust, deep-bodied Leporinus with a

stocky appearance. The mouth is terminal.

Counts. DR iilO; AR ii9; PR i14-i16; VR i8; LS 38-40; PDS 9-12;

CPS 16; TS 11-12; teeth 3-3 on premaxilla, middle incisors bifid, the

others strangely curved; 3-4 teeth on dentary, the last one tiny and

next to base of last big tooth.

Measurements. GBD 32-36% SL.

Pigmentation. The dorsum is grayish, and crossed by about 12-15

faint vertical bars and the ventrum is light tan. The sides are brown,

with at least two large, rounded black spots, the first spot under

dorsal fin base, the second just anterior to the adipose fin. Sometimes

there is an additional spot at the base of the caudal fin. The fins,

especially the anal, are dusky and heavily pigmented on the membranes.

The adipose fin is often edged in black.




























































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Distribution and Natural History

Range. It occurs in the Amazon and Orinoco basins, and the

Guianas.

Aoure distribution. Map: fig. 12. It is common throughout the

piedmont and llanos.

Habitat. It lives in a wide variety of streams and rivers from

both white and blackwater habitats.

Abundance. COMMON.

Number of specimens examined. 230 from 64 collections.

Food. OMNIVORE. It feeds mostly on aquatic plants, but seeds and

carrion (fish flesh) are also included in its diet (K. Winemiller pers.

com.).

Reproduction. Probable strategy: r2. This species migrates down-

stream shortly after the first rains of the season. Spawning probably

takes place in midriver, in a fashion similar to Prochilodus mariae.

Migrations. This fish is known to accompany Prochilodus mariae on

annual migrations from the lowland floodplain habitats upstream to

piedmont and montane streams at the beginning of the dry season, and

thence back downstream when the seasonal rains begin.

Importance. It is of minor importance as a commercial species.


Leporinus sp.

Fatlip Leporinus Mije

Fig. 13. Map fig. 14. Couplet 9a.

Comments. This may be a new species. Its plain coloration and

meristic characters don't agree well with any described species.





































Figure 13. Leporinus sp.







Description

Illustrations. Fig. 13.

Diagnosis. This fish is quite plain, usually brown when adult,

with a small black spot at the base of the tail (fig. 13). The lip is

much thicker and more gently curved than in the similar L. friderici.

It has a more slender body as well. The lateral-line scale count of 42

to 44 is higher than for most other Apure drainage Leporinus species, in

which the number usually does not exceed 40.

Size. It reaches at least 250 mm SL.

Counts. DR iilO; AR ii9; PR i14-i18; VR i8; LS 42-44; PDS 12-13;

CPS 16; TS 13-14.

Measurements. GBD 26-30% SL in specimens above 90 mm SL.

Pigmentation. The body in adults is plain brown with a small spot

at the base of the caudal fin. Smaller specimens may have two or three

spots on the sides.

Distribution and Natural History

Range. It is known from the Orinoco Basin.

Apure distribution. Map: fig. 14. It occurs throughout the pied-

mont and llanos.

Habitat. It lives in muddy (whitewater) streams and rivers.

Abundance. COMMON.

Number of specimens examined. 81 from 25 collections.

Food. OMNIVORE. It is probably mostly vegetarian, but undoubt-

edly includes some aquatic invertebrates in its diet.

Reproduction. Probable strategy: r2.

Migrations. It makes annual migration similar to those of

Prochilodus mariae, and often accompanies that species during the


































































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the onset of the dry season in October or November).

Importance. It is of only minor commercial importance.


Leporinus sp. "aguaro"

Leporinus Mije

Fig. 15. Map fig. 16. Couplet 11b.

Comments. This species is similar to the species I have identified

as L. friderici, and could turn out to be the "real" L. friderici. If

that is the case, the species I have identified as L. friderici is

probably new. The original descriptions don't provide adequate informa-

tion to make a certain determination. L. friderici was described from

Surinam, presumably from blackwater rivers typical of the Guyana Shield

and coastal drainages. Leporinus sp. "aguaro," is apparently restricted

to similar habitats in the Apure drainage.

Description

Illustrations. Fig. 15.

Diagnosis. This species can be distinguished by pigmentation

pattern (fig. 15), which consists of three to four large spots along the

lateral midline plus a faint row of smaller spots above them on the

upper sides. It has fewer lateral scales (39 vs 42-44) than Leporinus

sp., and a relatively larger eye (the eye diameter greater than half of

the interorbital width) than L. friderici (less than half the interor-

bital width).

Size. It grows to at least 200 mm SL.

Morphology. The snout length is about equal to the interorbital


width.






































Figure 15. Leporinus sp. "aguaro".







Counts. DR iilO; AR ii9; PR i15-i16; VR i8; LS 37-39; PDS 10-12;

CPS 16; TS 11-12, with 5-6 scales above and five below the lateral

line.

Measurements. GBD 29-32% SL; HL 28-31% SL; EYE in HL 22-34%, in

SNT 53-96%, in IO 54-86% (the large variation in eye diameter is due to

allometry, juveniles have relatively larger eyes).

Pigmentation. The body is brown, darkest on the dorsum and

becomes gradually whiter towards the belly. The sides are marked with

three or four large black rounded spots, the first under the posterior

dorsal-fin base, the second just anterior to the adipose fin, the third

under the adipose fin, and the fourth on the caudal peduncle. Another

series of seven smaller spots is situated above these on the upper

sides. Each scale has a dark spot on its margin, resulting in up to 10

horizontal rows of dots; in some specimens the dark pigment completely

outlines each scale, forming a net-like pattern.

Distribution and Natural History

Range. It occurs in the Orinoco Basin.

Apure distribution. Map: fig. 16. It is known only from the

Aguaro River system.

Habitat. It occurs in blackwater streams and rivers.

Abundance. RARE.

Number of specimens examined. 3 from 3 collections.

Food. OMNIVORE. It is probably mostly herbivorous.

Reproduction. Probable strategy: r2.

Migrations. They are probable, but not documented.

Importance. It is potentially valuable as an ornamental.

















































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Leporinus cf striatus Kner 1859

Striped Leporinus Mije Rayado

Fig. 17. Map fig. 18. Couplet 8a.

Specific Synonymy

Leporinus striatus Kner 1859:171 pl. 8, fig. 18 (type locality:

Orissanga, Rio Parana and Caicara Mato Grosso, Brazil); Pellegrin

1899:157 (Apure River, Venezuela); Gdry 1977:170 (key).

Ledorinus (error) striatus Bertoni 1939:54 (Paraguay).

Salmo tiririca Natterer in Kner 1859:172 (?based on same type as

L. striatus Kner).

Comments. Since L. striatus was described from southern Brazil, it is

doubtful that the same species is present in the Apure drainage. Fowler's

illustration (1950) depicts a fish with only two stripes on the side. I

have examined the holotype (FMNH 53366) of the similar L. arcus Eigenmann

1912, which was described from British Guiana, and which would perhaps be

more likely to occur in Venezuela. It is, however, a different-looking fish

with only four stripes on either side (vs five in Apure specimens). In

addition the dorsum in the L. arcus holotype is uniformly dark, and though

the pigment may have obscured a stripe that may have originally been

present, none is shown in the figure of the holotype (Eigenmann 1912:pl. 62,

fig. 3). The holotype has 36 lateral-line scales (vs 38 in Apure specimens)

and ten transverse scales (vs 12). Finally, specimens of the type series of

L. arcus greatly exceed the maximum recorded size for L. striatus. These

data strongly suggest that the Apure specimens are not L. arcus. I suspect

that the Apure drainage form is undescribed, but this cannot be finally

resolved until it can be compared with fresh material from the type locali-

ties of L. arcus and L. striatus.






































Figure 17. Leporinus cf striatus






Etymology. STRIATUS = striped.

Description

Illustrations. Fig. 17; Axelrod et al. 1971:F-378.00; RomAn

1983:99.

Diagnosis. The color pattern of black stripes over a yellowish

background (fig. 17) is unique for Apure drainage anostomids. In this

regard it is superficially similar to Anostomus ternetzi, but that

species has a superior mouth and is much more slender and elongate.

Size. It is seldom found over 100 mm SL, but can reach 175 mm.

Morphology. This is a robust, stocky fish. The horizontal diame-

ter of the orbit is longer than the snout length, but slightly less than

the interorbital width. The mouth is subterminal.

Counts. DR iilO; AR ii8; PR i14; VR i8; LS 38; PDS 10; CPS 16; TS 12;

teeth 3-3, 3-3 or 4-4 (if four the last very small), the teeth are curved

incisors with brown tips, not truly cuspid but with irregular margins. The

center of the roof of the mouth has a bulbous papillose projection.

Measurements. GBD about 29% SL; HL 36% SL; EYE 36% HL. IO 39% HL.

Pigmentation. Juveniles up to about 40 mm SL show vertical

banding instead of the typical adult pattern of horizontal light and

dark stripes. They also have a jet-black adipose fin with a light

center, a dark crescent along base of tail with darker dot at midpoint,

and a black anal fin. Juveniles also lack pigment on the fontanel and

have about three vertical stripes anterior to the dorsal fin. At 19 mm

SL one juvenile had only vertical barring, but another specimen of 37

mm SL had the beginnings of horizontal stripes.

Adults have a black line along the dorsal midline of the body in

front of the dorsal fin, but the dorsal midline of the head has a white




77

stripe. They have five additional dark stripes on either side, the

first of which extends from the occiput along the uppermost sides and

back to the adipose fin. The second extends from the snout, back above

the eye to the uppermost caudal-fin base. A third extends from the

snout through the eye across the opercle and back along the midside to

midtail. The fourth extends from the pectoral-fin base (which is dark

dorsally near its base) back to the anal fin and onto the lower caudal

peduncle. The fifth stripe is faint and extends from the lower pectoral-fin

base to the pelvic-fin base. The adipose and anal fins are black, the other

fins white, tan, or transparent.

Distribution and Natural History

Underwater observations confirm that this species usually travels

in schools, as do most Leporinus. However, since they occur over rocky

substrates where seines are easily avoided, most of our collections are

of very few specimens.

Range. It has been reported from the Orinoco, Amazon and Parana

basins.

Apure distribution. Map: fig. 18. It is most common in the Andean

piedmont and streams in the upper llanos.

Habitat. L. striatus is most commonly found in clear streams and

rivers, with fast currents and rocky substrates.

Abundance. COMMON.

Number of specimens examined. 109 from 35 collections.

Food. OMNIVORE. It is probably mostly herbivorous.

Reproduction. Probable strategy: r2.

Importance. Ornamental.

























































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Leporinus yophorus Eigenmann 1922

Banded Leporinus Mije Rayada

Fig. 19. Map fig. 20. Couplet 8a.

Specific synonymy.

Leporinus yophorus Eigenmann 1922:233 (type locality: Barrig6n,

Colombia, upper Rio Meta)

?Leporinus affinis Pellegrin 1899:157 (Apure River, Venez.).

Types: Holotype: CAS 61680 (formerly IU 15025).

Comments. There are many names available for vertically banded Lepori-

nus species. Most often they are called L. fasciatus, but L. latofasciatus

and L. pearsoni are other possibilities for Venezuelan specimens. The Apure

form agrees almost exactly with Eigenmann's (1922) original description of

L. vophorus. Since both the habitat and drainage coincide, I feel reasona-

bly confident about this identification. It is also possible that L. vopho-

rus could prove to be a junior synonym of one of the older names such as

Leporinus affinis GUnther 1864, which was described from Para, Brazil, and

was cited by Pellegrin (1899) from the Apure River. However, that species,

which has nine vertical bars according to Gdry (1977), is probably not

present in the Orinoco Basin.

Etymology. Y = the letter "Y" refers to the Y-shaped bar on the

side, OPHORUS = bearer of.

Description

Illustrations. Fig. 19; Eigenmann 1922:317 pl. 20, fig 4.

Diagnosis. The color pattern (fig. 19) is distinctive since this

is the only Apure drainage Leporinus with bold vertical bars.

Size. It grows to about 180 mm SL






































Figure 19. Leporinus yoDhorus.




82

Morphology. This is an elongate, slender Leporinus with a terminal

mouth.

Counts. DR iilO; AR ii9; PR i15-i16; VR i9; LS 42-45; PDS 14-16;

CPS 16; TS 12, 6 above, 5 below lateral line.

Measurements. Eigenmann (1922) gave the following measurements for

the type: Head 5.0 in SL, depth 4.66, eye 2 in snout, 4.5 in the head,

2.0 in the interorbital; isthmus 3.5 in head; depth of caudal peduncle

2.4 in head. Apure specimens: GBD 23% (adults) to 29% (juveniles) SL;

Eye 18% (adults) to 29% (juveniles) HL.

Pigmentation. The body is yellowish with seven wide vertical bars.

The second bar is divided at the level of the lateral line to form a "Y"

that extends dorsally.

Distribution and Natural History

Range. It occurs in the Orinoco Basin.

Apure distribution. Map: fig. 20. It is known only from four

localities, three from within or near the Apure River itself, and one

from the Tucupido River in the piedmont.

Habitat. There are insufficient numbers of collections to

permit a thorough characterization of this species' habitat.

The one adult captured was collected from a site in the Andean piedmont

in a rapidly flowing, rocky river. The juvenile specimens were taken

from near shore in muddy-bottomed side pools of the Apure River near

Bruzual, amidst aquatic vegetation and submerged grasses. Thus, it

seems that llaneran whitewater rivers are this species habitat.

Abundance. UNCOMMON, but it is probably difficult to capture.

Number of specimens examined. 84 from 11 collections.

Food. OMNIVORE. It is probably mostly herbivorous.





























































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Reproduction. Probable strategy: r2.

Importance. Ornamental.


Pseudanos gracilis (Kner) 1859

Banded Headstander Cabezibajo Rayada

Fig. 21. Map fig. 22. Couplet 4b.

Generic synonymy

Pseudanos Winterbottom 1980:24 (type species: Schizodon trimaculatus

Kner 1859:161, by original designation).

Specific synonymy

Schizodon gracilis Kner 1859:160 (type locality: Rio Guapor6, Brazil).

Anostomus gracilis Myers 1950:184; Kn6ppel 1972:268 (diet); Mago L.

1970:75.

Pseudanos qracilis Winterbottom 1980:24 (redescription).

Types. Holotype: NMW 57-119.

Description

Illustrations. Fig. 21; Winterbottom 1980:83 fig. 20.

Diagnosis. The minute, superior mouth distinguishes this species

from all Leporinus species. The pigmentation pattern (fig. 21) is also

characteristic, and useful to distinguish it from P. qracilis, and

Anostomus ternetzi.

Size. It grows to 160 mm SL, but most individuals are smaller.

Morphology. The body is fusiform but somewhat compressed.

Counts. DR iiil0; AR iii8; PR i13-i16; VR i8; LS 43-47; PDS 11-15;

CPS 16; TS 10, teeth 4-4 in both jaws (each with 2 to 4 cusps, though

these can wear down, and become less conspicuous).

Measurements. GBD 18.3-25.4% SL







































Figure 21. Pseudanos gracilis.






























































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Pigmentation. The pigmentation pattern varies in preserved material,

but the dorsum never has cross bars. The body has up to 11 thin horizontal

stripes formed by rows of dots (one per scale). A median predorsal stripe

is also present. The midlateral sides are marked with a dark stripe or with

series of up to four large rounded spots.

Etvmology. PSEUD = false, ANOS = the first four letters of Anostomus,

a very similar genus; GRACILIS = slender.

Distribution and Natural History

Range. It occurs in the Amazon and Orinoco basins.

ADure distribution. Map: fig. 22. It has been found only in the

Aguaro River system. Interestingly, the distribution of this species seems

to complement that of P. irinae, which is absent from the Aguaro systems but

present in suitable blackwater habitats of northern Apure and Barinas.

Habitat. It is restricted to blackwater streams with abundant aquatic

vegetation.

Abundance. RARE.

Number of specimens examined. 8 from 5 collections.

Food. OMNIVORE: It feeds on plant remains, fungi, algae, detritus,

sand and terrestrial insects (Winterbottom 1980).

Reproduction. Probable strategy: rl.

Importance. Ornamental.


Pseudanos irinae Winterbottom 1980

Three spotted Headstander Cabezibajo de Tres Puntos

Fig. 23. Map fig. 24. Couplet 3a.

Specific synonymy

Pseudanos irinae Winterbottom 1980:27 (type locality: Orinoco River

system, Cano de Quiribana where it empties into the Orinoco).






































Figure 23. Pseudanos irinae.







Anostomus trimaculatus (non Kner 1859) Eigenmann 1912:295 (Essequibo

River; Myers 1950:184 (upper Orinoco); Ramirez 1957:157 (Venez.);

Mago L. 1970:75 (Venez.).

Types. Holotype: CAS 58809. Paratypes: deposited at AMNH, BMNH,

CAS, FMNH (3) (Winterbottom 1980), catalog numbers not given.

Etymology. IRINAE = named for the wife of Dr. R. Winterbottom,

Irina.

Description

Illustrations. Fig. 23; Axelrod et al. 1971:F-41.00 (identified as

Anostomus trimaculatus); Winterbottom 1980:83, fig. 21.

Diagnosis. The pigmentation pattern is distinctive (fig. 23).

Size. It grows to about 100 mm SL.

Morphology. The body is fusiform and somewhat compressed.

Counts. DR 13-14; AR iii7-iii8; PR i13-i16; VR i7-i8; LS 41-45;

CPS 16.

Measurements. GBD 23-29% SL.

Pigmentation. This fish is yellow-brown, paler below, with the dorsum

crossed with irregular, thin vertical bars, and two to four dark spots or

blotches on the sides. The first of these spots covers scales three to five

of the lateral-line series and is situated partly beneath the lateral line),

the other spots cover scales 16-19, 29-30, and 41-46. The first and third

spots are often faint and may be absent. The area around each spot is

lighter.

Distribution and Natural History

Range. It occurs in the Orinoco and Essequibo basins, of Colombia,

Venezuela and Guyana.






























































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Apure distribution. Map: fig. 24. It is so far known only from the

SuripA River in Barinas, and the Guaritico system of northern Apure state.

Habitat. It is restricted to black or clearwater streams in the

savannas of the low llanos. and is usually found in aquatic vegetation.

Abundance. UNCOMMON.

Number of specimens examined. 17 from 6 collections.

Food. OMNIVORE. The diet of this species is probably similar to that

of the similar, preceding species which feeds on plant remains, fungi,

algae, detritus, and insects.

Reproduction. Probable strategy: rl.

Importance. Ornamental.


Schizodon isoqnathus Kner 1859

Boquimi, Coti, Pijotero Schizodon

Fig. 25. Map: fig. 26. Couplet 5a.

Generic synonymy

Schizodon Agassiz 1829:66 (type species: Curimatus fasciatus Agassiz, by

monotypy).

Specific synonymy

Schizodon isoqnathus Kner 1859:163 pl. 6 (not pl. 7 as stated in

the text; type locality: Rio Cujaba).

Anostomus isoqnathus Peters 1877:472 (San Fernando de Apure, Venez.).

Comments. Recent investigations by biologist Maria Esther Antonio

of the Universidad Central de Caracas indicate that this species is

distinct from the Amazonian S. isoqnathus, and perhaps new.

Etymology. SCHIZ = divided, ODON = teeth; ISO = even, equal,

GNATHUS = jaws.




Full Text

THE CHARACIFORM FISHES
OF THE APURE RIVER DRAINAGE, VENEZUELA
By
DONALD C. TAPHORN
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1990

ACKNOWLEDGMENTS
Saying thank you is hardly adequate to repay the many people who have
helped me in this project. They have given me tremendous gifts: precious
time, sound advice and useful suggestions, as well as moral and financial
support and many other things. They helped get the fish out of the water
and into a jar in the museum, get the sometimes jumbled ideas in my head
into some semblance of order, get my body out of bed and into the lab, and
get the words out of the computer and onto this page. But thank them I
must and I do so sincerely.
My parents and family are first in this list of special people who
have helped. They are always with me in spite of the large distances that
have separated us. Next I thank the many friends, professors, colleagues
and students who have helped to make this dissertation possible. They are
Dr. Carter Gilbert, cochairman of my committee for these many years and
tireless in the dreary task of wading through and correcting the rough
drafts, Drs. Horst Schwassmann, Frank Nordlie, and Nigel Smith, who pa¬
tiently served on my committee and suggested many helpful improvements and
Dr. Craig Lilyestrom, his wife Maria and their children Lynda and Wayne who
shared a home with me in Venezuela. During my visits to Gainesville, I
enjoyed the warm hospitality of Dr. and Mrs. Carter Gilbert, Mr. and Mrs.
George Burgess, and Mr. and Mrs. Terry Converse.
Dr. Kirk Winemiller, Mr. Leo Nico and Mr. Stewart Reid worked on
postgraduate degrees in our laboratory, shared in the field work, and

generously provided specimens and information on the ecology of Apure drain¬
age fishes from their master’s and doctoral work.
I also thank the friends and colleagues who helped pull the other
(usually the deep) end of the seine, Craig Lilyestrom, Craig and Carmen
Olds, Leo Nico, Stewart Reid, Kirk Winemiller, Oscar Leon M., Naboth
Mont ilia A., Mike Taphorn, Pablo Osman, Larry Page, Carter Gilbert,
Richard Franz, Jack Karr, Jamie Thomerson, Guillermo and Ramón Feo,
Alex Flecker and many others too numerous to mention here. Special
thanks go to Ing. RNR Angelina Licata and Mr. Oscar Leon Mata for the
drawings and to Lie. Jairo Pérez and Ing. Aniello Barbarino D. for the
maps. Three technicians have worked with me in the Fish Collection,
Aldo García, Eric Sutton and Keyla Marchetto; I am greatly indebted to
them for their assistance with the unending museum chores of processing
and cataloging the specimens, preparing labels and card files, and
keeping the computer records up to date. Several of my students in the
Environmental Engineering Department have done thesis work on fishes,
and many also held student jobs in the Fish Collection and helped with
the museum chores. Special thanks go to Pablo Osman, Norberto Saavedra,
Enrique Marzola, Aniello Barbarino Duque, Cecelia Gómez, Guillermo
Cedeño, Ricardo Smith, Yury Hernández. I also thank Dr. Richard Schar-
gel for his helpful talks about Apure drainage soils and Prof. Heberto
Pacheco and the other personnel of the Cartographic Center of UNELLEZ
who helped obtain topographical maps.
The Secretario de Investigación de la UNELLEZ, FONAIAP, FUNDA-
CITE, CONICIT all provided financial support for this project.

TABLE OF CONTENTS
page
ACKNOWLEDGMENTS ii
ABSTRACT v
INTRODUCTION 1
MATERIALS AND METHODS 14
Collecting 14
Curation 14
Material Examined 14
Format 15
RESULTS 21
Characiformes 23
Anostomidae 31
Characidae 100
Characidiidae 606
Chi 1 odontidae 664
Ctenoluciidae 670
Curimatidae 677
Cynodontidae 721
Erythrinidae 739
Gasteropelecidae 760
Hemiodontidae 766
Lebiasinidae 777
Parodontidae 796
Prochilodontidae 808
DISCUSSION 823
Ecological Considerations 823
Zoogeographical Considerations 854
LITERATURE CITED 864
APPENDIX 887
BIOGRAPHICAL SKETCH 891
i v

Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
THE CHARACIFORM FISHES
OF THE APURE RIVER DRAINAGE, VENEZUELA
By
DONALD C. TAPHORN
May, 1990
Chairman: Dr. Carter R. Gilbert
Major Department: Zoology
The Apure River drainage of Venezuela collects waters from many
different aquatic habitats that vary greatly in geology, climate, vegeta¬
tion and altitude. Over ten years of sampling has yielded 354 species of
freshwater fishes pertaining to ten orders and forty-one families. This
report includes taxonomic synonymies, keys, illustrations, morphological
descriptions, distribution maps, data on abundance, habitat, life history
strategy and diet for 138 Characiform species of the families Anostomidae,
Characidae, Characidiidae, Chilodontidae, Ctenoluciidae, Curimatidae,
Cynodontidae, Erythrinidae, Gasteropelecidae, Hemiodontidae, Lebiasinidae,
Parodontidae, and Prochilodontidae.
Fish distribution within the Apure River drainage is primarily gov¬
erned by two parameters, altitude and water chemistry. The few characi-
forms adapted to mountain and piedmont conditions tend to be restricted
in distribution to only those areas.
v

In the llanos, two assemblages are recognized. Whitewater, muddy
streams are the most common habitat in the drainage, but few characiforms
are restricted only to them. Blackwater habitats with distinct fish assem¬
blages exist in isolated pockets in the Aguaro-Guariquito drainage, in the
Caicara Creek drainage of northern Apure state, and in west-central Barinas
state.
The drastic changes in climate between wet and dry season in the
Apure River drainage have molded characiform life history strategies. Most
characiforms have adopted an rl or r2 life history strategy. Of 68 species
that have adopted the rl strategy, most are small omnivorous fishes with
low juvenile and adult survivorship, low fecundity per reproductive bout
(but repeated reproduction throughout the rainy season), short generation
times (many live less than one year), and widely fluctuating population
densities during the year. The other 70 species present have been classi¬
fied as r2 strategists. These species typically are larger, are often
migratory, and reproduce annually in a single massive spawning event.
Juvenile survivorship is low, but adults live for several years and reach
larger sizes. Populations fluctuate greatly from wet to dry season because
of the high juvenile mortality. No characiforms in the Apure drainage have
adapted the K-strategy.
The Apure drainage ichthyofauna is a mosaic of relatively young
species that have evolved in the Whitewater habitats and blackwater species
probably derived from an older Guyana Shield fish fauna.
vi

INTRODUCTION
Tropical aquatic ecosystems are suffering rapid modification and
deterioration as man’s growing population makes ever-increasing demands for
water, forest products, and agricultural land. Many Venezuelan rivers are
already so polluted that we will never know much, if anything, about their
original fish faunas. Fishes are suffering loss of habitat to wetlands
drainage programs, habitat modification by dams and dikes, pollution from
urban sewage, agricultural fertilizers and biocides sedimentation from
deforestation and poor upland watershed management, overfishing, and the
introduction of exotic species.
Efforts to produce effective management plans for a rational develop¬
ment of tropical aquatic resources suffer from an almost total lack of basic
information about the aquatic organisms that occupy these systems. Not even
lists of the fishes, mollusks, crustaceans, aquatic insects, plants, etc.,
can be found for most tropical rivers, much less data on abundance, life
histories, critical habitat requirements or other aspects of their biology.
Furthermore, the majority of these organisms have never been sufficiently
sampled, nor has their taxonomy been adequately studied. The literature
provides few guides to the identification, and for the most part, original
species descriptions are hidden in old primary references not usually avail¬
able to workers in South America. What little information is available is
seldom written in Spanish, and so is of little use to management officials.
For these and many other reasons, inventories of tropical aquatic organisms
1

2
are desperately needed. The decision to undertake a faunal survey of the
Apure River drainage, part of which forms the basis of my doctoral disserta¬
tion, is in response to this need.
In South America the Orinoco River Basin (fig. 1) collects the waters
of a vast area covering approximately 1,123,000 km^, second only (although
the Rio La Plata is longer) to the immense Amazon River Basin to the south,
which drains some 7,000,000 km^ (Zinck 1977, Smith 1981). About three-
fourths of the area of the Orinoco Basin falls under Venezuelan jurisdic¬
tion, whereas the rest is Colombian. The river traverses some 2,148 km in
its journey to the sea from its currently recognized source near Cerro
Delgado-Chalbaud in the ancient mountains of the Guyana Shield near the
border with Brazil. In my opinion, the true source of the Orinoco is the
Guaviare River, which originates in the Andes of Colombia and contributes
much more water than the Ventuari. In any case, the Orinoco flows west from
its Venezuelan source, at first descending through dense tropical forest
that grows in the wel1-weathered rocky drainages of the Guyana highlands.
After sharing a portion of its reddish-brown waters with the Amazon via the
Casiquiare Channel, it receives its first major tributary, the Ventuari, and
then veers north to run along the northwestern edge of the Guyana Shield
where it marks the boundary between Colombia and Venezuela. Several major
tributaries enter the Orinoco from Colombia in this stretch (fig. 1), among
the largest of which are the Guaviare, Vichada, Tomo and Meta. At Puerto
Páez, the Orinoco turns slightly to the east, and receives the black waters
of the Cinaruco and Capanaparo rivers that drain the grassland savannas of
southern Apure state. Rocks of the Guyana Shield still project though sandy
bottom sediments at this stage. Gradually, the river bends more eastward.
Just before reaching Caicara, the muddy waters of the Apure River dump

Figure 1. Orinoco River Basin in Venezuela.


5
their Andean sediments into the main channel of the Orinoco. Since the
Orinoco is often full during peak flood of the Apure, it tends to hold back
the waters of the Apure, and has formed an interior delta where vast areas
flood each year. The Orinoco changes course at this point to run in a
northeasterly direction towards the sea. It is bordered by the semi-arid
central plains to the north, and the humid Guyana Shield to the south. It
picks up vast quantities of black waters from the Caura and Caroni rivers
before it enters its highly productive delta and flows into the Atlantic
Ocean just south of the island of Trinidad. The average annual outflow is
about 33,000 m^/s (Zinck 1977). The Amazon, by way of comparison, delivers
an annual average of about 175,000 m^/s (Smith 1981).
The Apure River drainage lies mostly within 7° and 10° North lati¬
tudes, and 66° and 72° West longitudes. It is Venezuela’s largest tributary
(in terms of drainage area) of the Orinoco (fig. 2a & 2b) and drains 167,000
km2, or nearly 15% of the total area of the Orinoco Basin. Its average flow
of nearly 2000 m^/s (Zinck 1977) divided by the drainage area gives a figure
of about 378 mm/year runoff, which is about 24% of the mean precipitation
(Saunders & Lewis 1988a). It is exceeded in Venezuela by only two other
tributaries, both blackwater rivers from the nutrient poor soils of the
Guyana Shield, where rainfall is much higher. The Caroni River, which
drains some 93,000 km2 delivered over 4,000 mVs before the enormous Raúl
Leone (Guri) Dam was built. The Caura River, slated for future hydroelec¬
tric development, carries 2,700 m^/s. The Guaviare River which drains
159,500 km2 and the Meta River with 103,000 km2 are two of the major Orinoco
tributaries in Colombia (Zinck 1977). The headwaters of the Apure drain the
southeastern flank of the Andes, and are located approximately 800 km by
river from the Orinoco River. Over 15% of the drainage is mountainous

Figure 2a. Apure River Drainage in Venezuela.


Figure 2b. Apure River Drainage in Venezuela showing collecting sites.


10
(above 500m in elevation). Transit time from the Andes to the Orinoco
varies seasonally in the range of one to four weeks (Saunders & Lewis
1988a,b). On the way, the numerous branches of anastomosing rivers cross
the llanos, flat alluvial plains that formed in the late Pleistocene over
older marine sediments. In the wet season, most of the lower part of the
drainage is inundated by sheet flooding. The floodplain of the Apure is
much more extensive than that of most rivers. An internal delta of about
5000 km^ forms where the Apure and the Arauca (the first major tributary of
the Orinoco to the south) flow into the Orinoco. When discharge is high in
the Orinoco, a large inland sea forms between the mouths of the Capanaparo
river (the next Orinoco tributary to the south) and the Apure. The alter¬
nation of wet and dry seasons causes annual changes in river level of about
six to eight meters. Using river level values from San Fernando de Apure,
Saunders & Lewis (1988a) defined four hydrologic phases to describe the
relationship between the river and the floodplain. The river is not in
contact with the floodplain during the first, low-water phase, which corre¬
sponds to stage heights below 38 m. The rains, which usually start in
April or May, initiate the second, rising-water phase, during which the
river is still within its banks (38-42m). The inundation phase commences
when stage heights surpass 42m, and river water floods onto the surrounding
floodplain. Flooding usually begins in July, and peaks in September or
October. When the dry season sets in and water levels drop to below 42m,
the fourth, or falling-water phase begins (42-38m).
Of all of the Orinoco tributaries, the Apure is by far the most acces¬
sible, with good, all-season roads crossing all of its major drainage sys¬
tems. This factor alone was enough to select it as my study site, but in
addition to easy access the climate and topography are extremely

11
diverse. As a result, many different aquatic biotopes are included in the
drainage. There are cold mountain streams that tear through the Andes
at breakneck speeds; tepid, blackwater lowland swamps; large "white-
water" muddy rivers that meander slowly through the lower llanos (a
Venezuelan term for plains or savanna); and all the varied intergrada¬
tions of these that can be imagined. The good roads in this region owe
their existence to the fertile soils and resultant extensive agriculture
that has developed. Deforestation has proceeded virtually unchecked
since colonial times. As more and more forests give way to pastures and
crops, the aquatic fauna becomes increasingly degraded. Massive fish
kills occur almost as a matter of routine when the first rains fall
after the months of drought and wash the accumulated pesticides and
fertilizers into the drainage in one large dose. Natural aquatic habi¬
tats are disappearing rapidly and fish populations are suffering accord¬
ingly. This too gave the Apure drainage a high priority as a study
area, inasmuch as the fishes sensitive to these changes may not be
around indefinitely.
Another factor in the selection of the Apure drainage is that the
Apure River and its larger tributaries support a large commercial fishery,
and provide a significant source of protein and extra income for rural
families. Almost no basic biological data is available for most species.
Declining catches in recent years and local conflicts over the best use of
aquatic resources in the region has local and federal governments concerned.
Most commercial species are migratory, moving in tune with the drastic
seasonal changes in water level and flow because of the accentuated wet
and dry seasons typical of the western llanos. Funding for impact
studies of local dams on commercial species helped pay for the sampling.

12
The Andean piedmont was characterized as an area of recent speciation
by Mago L. (1978), but Andean piedmont rivers, flowing out through narrow
valleys into the plains, offer excellent opportunities for hydroelectric
power generation. The Venezuelan government, eager to diversify its energy
resources now that the end of their vast petroleum wealth is in sight, has
projects for dams on almost every major piedmont river system. These dams
have denied the commercial species, particularly the coporo, Prochi 1odus
mariae, access to the upland piedmont and montane lotic systems, and disrupt
the annual cycle of flood and drought used by migratory species as the cue
to make their moves. Commercial species populations are on the decline
throughout the Apure drainage, but direct causes are not easily identified
because of the wide array of concurrent negative impacts that are occurring.
As the commercial fish catch drops, concern is mounting, and governmental
interest in finding a solution is growing. But again, lack of basic infor¬
mation about fish populations and ecology hinders progress.
In his seminal work, "Lista de los Peces de Venezuela incluyendo un
Estudio Preliminar sobre la Ictiogeografía del País," Mago Léccia (1970)
lamented the deplorable state of knowledge of the Venezuelan freshwater fish
fauna. He noted that most of the pioneering ichthyological works of Eigen-
mann, Steindachner, Regan, Kner, Pellegrin, Norman, Fowler and others were
based on specimens from just about everywhere in South America except Vene¬
zuela. Only the Maracaibo Basin, studied in detail by Leonard P. Schultz
(1944a & b, 1949), was relatively well known. Since then, the pace of
ichthyological research in Venezuela has quickened considerably. This is
principally due to the work of Dr. Mago L. and his students, but contribu¬
tions have also been made by Agustín Fernández Yépez, Felipe Martín and
Manuel Ramírez. However, the present state of knowledge on the freshwater

13
fishes of Venezuela is far from complete, and most areas still need to be
better sampled.
Neotropical characiform fishes have received considerable attention
from ichthyologists. C. H. Eigenmann provided the basis for most modern
studies with a series of papers published in the late 1800’s and early
1900’s that culminated in several huge tomes such as his American Characidae
series. Other workers such as E. Ahí, J. Bóhlke, H. Fowler, J. Géry, J.
Hoedeman, R. von Ihering, W. Ladiges, K. Lüling, S. Meek, H. Meinken, N.
Menézes, G. Myers, A. and R. Ribeiro de Miranda, T. Roberts, H. Travassos,
S. Weitzman and R. Vari have continued the effort to give us a much better
idea of the tremendous diversity of form and function represented in this
group.

MATERIALS AND METHODS
Collecting
Many different collecting methods were employed, but most collections
were made with small, fine-meshed seines. Gill nets (both multifiber nylon,
and monofilament), beach seines, cast nets (both fine and commercial mesh
sizes), hand nets, and small otter trawls (in the Apure River’s main chan¬
nel ) were also used. We also fished with electrofishing gear, rotenone, and
hook and line. Some specimens were obtained from commercial fishermen.
Fishing effort was not vigorously controlled at each site. Usually
sampling continued until no new species appeared. From 1977 to 1989 we
made a total of 875 collections in the Apure drainage (fig. 2b).
Curation
Specimens were preserved in 10-15% formalin solution, and later trans¬
ferred to either 50% isopropanol, or 70% ethanol. Larger specimens were
usually slit on the right side to ensure complete preservation.
Material Examined
Most of the specimens examined as part of this study have been depos¬
ited in the Museo de Ciencias Naturales de la UNELLEZ--Guanare (MCNG).
Additional material is deposited in the Field Museum of Natural History in
Chicago (FMNH), the museum of the Illinois Natural History Survey (INHS),
the Museo de Biología de la Universidad Central de Venezuela in Caracas
(MBUCV), the museum of Sacramento State College, in Sacramento California
(SSC), the Texas Memorial Museum in Austin (TMM), the Florida Museum of
14

15
Natural History in Gainesville (UF), and the United States National Museum
(USNM) in Washington. Other museum acronyms used in this study are:
AMNH--American Museum of Natural History; ANSP--Academy of Natural Sciences,
Philadelphia; BMNH--British Museum of Natural History; CAS--California
Academy of Sciences; CM--Carnegie Museum (now mostly in FMNH and CAS);
IU--Indiana University (these specimens are now located mostly in FMNH or
CAS); MCZ--the Museum of Comparative Zoology at Harvard; MNHN--Muséum
National d’Histoire Naturelle, Paris; MZUSP--the Museum of Zoology of
the Sao Paulo University in Brazil; NMW--the Vienna Natural History
Museum; SU--Stanford University (now at CAS, and designated CAS-SU);
ZMA--Zoologisch Museum of Amsterdam; ZMB--Zoologisch Museum of Berlin.
Format
The following format has been used to organize the information pre¬
sented here. A brief introduction summarizing pertinent data for each
family is followed by a key to the species known to be present in the Apure
drainage. These sections are followed by the species accounts, which are
arranged, in sequence, under the following headings:
1. Genus--species--author--year (centered at top).
2. Common Name in Spanish & Common Name in English (if known).
3. Fig. #. This is the figure number(s) of the illustration(s).
4. Map: fig. #. This is the figure number of the dot map showing the
Apure drainage distribution.
5. Couplet. #. The couplet(s) where the species keys out is given.
6. Generic synonymy. This section, given as part of the species account
for the first species of each genus, includes the principal elements of a
generic synonymy: original name, describer, date, and type species. Only
primary junior synonyms are listed.

16
7. Specific Synonymy. The specific synonymy includes the original name,
describer, date, and page number, as well as the type locality (only given
for citations of original descriptions), junior synonyms, and selected
references to Venezuelan specimens or other important reports. It is not
intended as a complete synonymy, but in many cases it may be. In most cases
synonymies have been copied from the literature. When the original descrip¬
tion has been seen by me (many are in unavailable literature) this is so
indicated (as: seen).
8. Types. If readily available, museum location of the type material is
presented.
9. Comments. Comments are given on the taxonomic status of the species.
A synopsis of recent opinions on the status of the species or an explanation
of the validity of the name used is often presented.
10. Etymoloqy. The semantic roots or origin of the scientific name
are explained.
11. Description Usually just a subheading is given here.
12. Illustrations. All original illustrations in this report are listed
first, followed by references to selected illustrations of the species
available in the literature. The illustrations are not intended to portray
detailed anatomy (for example fin-ray counts) of the species, but rather
give an overall impression of the species’ basic morphology.
13. Di agnosis. A brief diagnosis of each species is given.
14. Size. The maximum size, as well as the size range of specimens usually
encountered are given.
15. Morphology. This section provides a brief physical description of
the most notable anatomical features of each species that are not mentioned
in the diagnosis.

17
16. Counts. Fin ray and scale counts are usually given. In most cases
only the range is given, but sometimes detailed counts are presented as the
count followed by the number of fish counted; for example: AR: 22(9) means
a count of 22 anal-fin rays was obtained for nine specimens examined.
The following abbreviations are used: DR--Dorsal Rays; AR--Anal Rays;
PR--Pectoral Rays; VR--Ventral or Pelvic Rays; LS--Lateral Scales;
LLS--Lateral-Line Scales; PDS--Predorsal Scales; TS--Transverse Scales,
(this is sometimes divided into scales above and below the lateral line;
CPS--circumpeduncular scale count; GR--gill rakers (if not indicated as
the total count of both upper and lower limbs, the figure given is the
count of the rakers found on the lower limb of the outer gill arch).
17. Measurements. Common body measurements are presented. The following
abbreviations are used: SL--Standard Length; TL--Total Length; GBD--Great-
est Body Depth; PDL--Predorsal Length; HL--Head Length; EYE--Eye Diameter;
SNT--Snout Length; IO--Interorbital Distance.
18. Pigmentation. Life colors are usually given along with pigmentation
patterns that remain in preserved specimens.
19. Pistribution and Natural History Comments that don’t fit into other
sections are given here.
20. Range. The overall geographic range is given.
21. Apure Pistribution. Map: fig. #. The number of the distribution map
for each species is given. In these maps, the collection localities are
plotted on a base map of the Apure drainage. Map scale sometimes required
that more than one locality be represented by a single dot. Brief comments
on the distribution within the Apure drainage are usually given. One should
bear in mind that dot maps of species distribution do not take into account
the fact that a few species in the Apure drainage are migratory, and thus

18
may be present in some areas during only part of the year. Also, non-
migratory species will spread out onto the floodplain in the wet season.
22. Habitat. A brief summary of the types of water bodies occupied by
the species is given.
23. Abundance. This is a somewhat subjective classification scheme, since
collection effort was not standardized to allow actual abundance data to be
calculated. I plotted the number of characid species versus the number of
collections, then arbitrarily drew separations between four categories as
follows:
RARE: If found in 1 to 5 collections.
UNCOMMON: If found in 6-25 collections.
COMMON: If found in 25-200 collections.
ABUNDANT: If found in more than 200 collections.
As indicated, these data are not quantitative, since fishing effort
varied considerably from site to site and not all specimens of each species
were preserved (especially true of the larger, more common species). Also,
the number of collections may include repeats from the same site. However,
the results are useful as a first approximation, and when used in conjunc¬
tion with the distribution maps they usually reflect a fairly accurate
picture of relative abundance, or at least ease of capture.
24. Number of specimens examined. This is the total number of specimens
collected in the Apure drainage as part of this investigation. It does not
include specimens examined from other collections.
25. Food. Species are usually classified as CARNIVORE, HERBIVORE,
OMNIVORE or DETRITIVORE, and the major items that comprise each species
diet are listed if known. This section is based on my own field obser¬
vations of live fishes and from stomach content analyses reported in

19
the literature, or done by myself, Dr. Craig Lilyestrom, Dr. Kirk
Winemiller or Mr. Leo Nico in our lab at UNELLEZ in Guanare.
26. Reproduction. Most of these data are from the literature, or
from data gathered by Dr. K. Winemiller for his doctoral research and
that was briefly summarized by Winemiller & Taphorn (1989). I use the
terminology introduced in that paper to classify the reproductive
strategy of each species:
"K-strategy" is characterized by high juvenile and adult survi¬
vorship, low fecundity, a long life and generation time (the time from
birth or hatching to reproductive maturity), iteroparity (spawning
several times per season) and a relatively stable population density
throughout the year.
"rl-strategy" is characterized by low juvenile and adult survi¬
vorship, low fecundity per individual but repeated spawning bouts
(iteroparity), a short life and generation time, and variable popula¬
tion densities throughout the year.
"r2-strategy" is characterized by low juvenile but high adult
survivorship, a high fecundity, a long life, semelparity (spawning all
at once, once per season) and by great fluctuations in the population
density.
27. Migrations. There are no comprehensive studies of migration of the
fishes in the Apure Drainage, but we are currently investigating the migra¬
tion and life cycle of the coporo, Prochilodus mariae (Barbarino D. &
Taphorn 1989). Local fishermen also provided information.
28. Importance. The commercial, sport, and ornamental uses as well as
the harmful or dangerous aspects of each species are discussed in this
section.

20
If no information is available on a given topic, that section is
omitted from the species account.
An abbreviated list of the material examined (the number of lots
or jars and the total number of specimens) is given for each species in
the Appendix. A more detailed list of specimens examined and locali¬
ties is available from the author. All specimens are deposited in the
Collection of Fishes, Zoological Musuem, Museo de Ciencias Naturales de
Guanare, at UNELLEZ, in Guanare, Venezuela.
Each family is presented in a separate section, and appears in
alphabetical order. A brief diagnosis of the family, with comments on
the number of species and their general ecology is followed by a key to
the species of the Apure drainage and the individual species accounts.
In some keys additional information useful for identification, follows
the species name in brackets.

RESULTS
After many years of sampling (1977-1989), during which some 875
collections were made in the Apure drainage, over 14,000 lots of specimens
have been processed, cataloged and identified. I have identified 354
species of fishes, belonging to 41 families (Table 1). The so-called
ostariophysans strongly dominate the fauna, and comprise 305 (86%) of the
354 species. The ostariophysans have the anteriormost four or five verte¬
brae fused and include, among South American groups, the orders Characi-
formes, Gymnotiformes and Siluriformes. The vast number of fishes col¬
lected and data generated by the project have made it necessary to limit
this dissertation to a subset of the total fish fauna. The order Characi-
formes was selected because of its great diversity, and a somewhat larger
volume of literature that is available for identification of the species.
This order includes 13 (32%) of the 41 families and 138 (39%) of the
species (Table 1). Only the order Siluriformes, with about 137 species,
matches the characiform fishes in diversity in the Apure drainage. The
taxonomy of South American fishes is still relatively poorly known, and
has been likened to the situation in North America 100 years ago. As more
collecting is done, and taxonomic revision continues, the total number of
species per family will continue to change, but the relative numbers of
species per family will probably not vary as greatly, except perhaps in
the order Gymnotiformes, which will probably outstrip the others in terms
of new species described.
21

22
Table 1. Composition of the Apure River Drainage Fishes.
Order
Family
Species
per
Family
Order
% of Total
1 RAJ I FORMES
1 Potamotrygonidae
4
4
1.1
2 CLUPEIFORMES
2 Clupeidae
2
5
1.4
3 Engraulidae
3
3 SALMONIFORMES*
4 Salmonidae
1
1
0.2
4 CHARACI FORMES
5 Anostomidae
12
138
38.9
6 Characidae
89
7 Characidiidae
11
8 Chi 1odontidae
1
9 Ctenoluciidae
1
10 Curimatidae
8
11 Cynodontidae
3
12 Erythrinidae
3
13 Gasteropelecidae
1
14 Hemiodontidae
2
15 Lebiasinidae
3
16 Parodontidae
2
17 Prochilodontidae
2
5 SILURIFORMES
18 Ageneiosidae
3
137
38.9
19 Aspredinidae
7
20 Astroblepidae
1
21 Auchenipteridae
10
22 Callichthyidae
7
23 Cetopsidae
3
24 Doradidae
16
25 Hypophthalmidae
1
26 Loricariidae
41
27 Pimelodidae
39
28 Trichomycteridae
9
6 GYMNOTIFORMES
29 Apteronotidae
16
31
8.7
30 Electrophoridae
1
31 Gymnotidae
1
32 Hypopomidae
3
33 Rhamphichthyidae
3
34 Sternopygidae
7
7 ATHERINIFORMES
35 Belonidae
2
10
3.0
36 Rivulidae
6
37 Poeciliidae
2
8 SYNBRANCHIFORMES
38 Synbranchidae
1
1
0.2
9 PERCIFORMES
39 Cichlidae
23
25
7.0
40 Sciaenidae
2
10 PLEURONECTIFORMES
41 Soleidae
2
2
0.6

23
Characiformes
Characiform fishes are ostariophysans, the group of fishes that
dominates freshwater habitats in most of the world. Ostariophysans are
combined into a group because they all have the first few vertebra modi¬
fied and linked to the surface of their bodies by a series of modified
bones to form what is presumed to be a listening device, or vibration
detector. Characiforms have gone farther with this device than most of
their cousins the catfishes, knifefishes, carps and suckers (the other
ostariophysans). They have developed a complete Weberian apparatus in
which the tripus is connected to the body of the third vertebra by means
of a vibrating lamina. This mechanism presumably transmits sounds or
vibrations picked up on the gas bladder to the inner ear (Géry 1977).
Better hearing could perhaps account in part for the dominance of the
ostariophysans in most freshwater ecosystems of the world (Fink & Fink
1981). Weitzman (1954, 1962) provided an extensive osteological defini¬
tion of characiform fishes and Fink & Fink (1981) discussed their rela¬
tionships with the other ostariophysans. The group is so diverse morpho¬
logically that they tend to defy succinct definition. About all one can
say is that they all have scales and teeth on the jaws or lips! Actually,
not even that is true of all characiforms since a few species from the
high Andes in southern South America have lost their scales completely and
the Curimatidae are toothless (at least as adults; juveniles have teeth in
the early developmental stages). Many are small schooling species that
have a chemoreceptory sensory system that, upon detection of substances
released by school members under attack or stress, will trigger a "fright
response," causing the school to adapt defensive maneuvers.

24
No consensus has yet been reached by workers on characiform upper-
level taxonomy. For many years almost all characoids were lumped in the
family Characidae (Eigenmann 1917, Weitzman 1962). The group was subdivided
by Greenwood et al. (1966), which (for Characiformes) is based largely on
the findings of Dr. Stanley Weitzman. Since then, the group, and the divid¬
ing lines between the families and subfamil ies, have been topics of constant
investigation, and debate. Géry (1977) proposed several modifications. In
Table 2 the two schemes are contrasted for South American groups.
Table 2. Comparison of family level taxonomy of Characiformes.
Greenwood et al.
Géry
Used Here
(12 families)
(11 families)
(13 families)
Characidae
Characidae
Characidae
Characidae
Characidiidae
Characidiidae
Characidae
Serrasalmidae
Characidae
Characidae
Crenuchidae
Characidae
Erythrinidae
Erythrinidae
Erythrinidae
Ctenoluciidae
Ctenoluciidae
Ctenoluciidae
Cynodontidae
Characidae
Cynodontidae
Lebiasinidae
Lebiasinidae
Lebiasinidae
Parodontidae
Hemiodidae
Parodontidae
Gasteropelecidae
Gasteropelecidae
Gasteropelecidae
Prochilodontidae
Curimatidae
Prochilodontidae
Curimatidae
Curimatidae
Curimatidae
Anostomidae
Anostomidae
Anostomidae
Hemiodontidae
Hemiodidae
Hemiodontidae
Chilodontidae
Curimatidae
Chilodontidae

25
Thus, the scheme of Greenwood et al. (1966) would leave South America
with 12 families of Characiformes to Géry’s 11. In more recent works, some
of these families have been promoted to familial or demoted to subfamilial
rank. Weitzman & Géry (1981) defined the Characidiinae. Nelson (1984)
listed Cynodontinae (which he earlier [1976] recognized as a family), as a
subfamily of Characidae; Vari (1989) treated it as a tribe of the Characi-
dae. Other groups have had the familial limitations better clarified and
justified, as for example Vari’s (1983, 1989) treatment of Curimatidae,
Prochilodontidae, Anostomidae and Chilodontidae. Nelson (1984) included
only eight families: Hemiodontidae, Curimatidae, Anostomidae, Erythrini-
dae, Lebiasinidae, Gasteropelecidae, Ctenoluciidae and Characidae, thus
demoting again the Prochilodontidae, Cynodontidae, Parodontidae and
Chilodontidae. Changes will continue. Except for the recognition of the
family Characidiidae, I have followed the classification of Greenwood et
al., for a total of 13 families. The correct spellings of Hemiodontidae
and Chilodontidae (vs Hemiodidae and Chilodidae) are also as yet unre¬
solved. The type genera are Hemiodus and Chilodus.
I have adopted this system as a matter of practical convenience. This
should not be interpreted as support for any particular position in the
controversy of higher level relationships of characiforms. For convenience,
and since taxonomic affinities are as yet unresolved at various levels, the
families, genera and species are discussed in alphabetic rather than taxo¬
nomic order.
The characiform fishes exhibit an incredible diversity of form
and occupy a wide array of ecological niches. Eigenmann (1917), Weitz¬
man (1962) and Fink & Fink (1981) all marveled at the incredible explo¬
sive radiation of this group in South America, which surpasses that of

26
the marsupials in Australia. Although many families contain widely
diverging morphotypes, I will try to briefly characterize each.
The Characidae comprises the most morphologically diverse group and
also includes the largest number of species. Although most species are
small minnow-like tetras ("sardinas" in Venezuela), this family includes
the piranhas, giant pacus, pike characins, strange tusked scale-eaters,
tiny killyfish-like inhabitants of blackwater streams, neon tetras, silver
dollars and palometas, and South American characid versions of trout,
salmon, clupeids and anchovies. The Erythrinidae is a small family of
large to medium fishes that are cylindrical, bowfin-like predators, typi¬
cal of shallow lentic habitats. The Ctenoluciidae is represented by only
a few species of elongate gar-like surface predators. Cynodontids are
large, riverine predators with keeled, compressed bodies, large eyes and
expanded pectoral fins adapted to life near the surface in strong cur¬
rents. Their extremely large canines give these piscivores a particularly
fierce appearance. Lebiasinids include two divergent groups: the "Creole
carps," Lebiasina, are medium-sized predators that inhabit mountain
streams, whereas Pvrrhulina and relatives are small, colorful fishes
typical of vegetation-choked lowland ponds. The Characidiidae and Paro-
dontidae include ecological equivalents of North American darters, adapted
to life in the riffles or mountain streams. The hatchetfishes of the
family Gasteropelecidae are bizarre, insectivorous, surface-dwelling
fishes with incredibly expanded coracoid bones (Weitzman 1954, 1960), and
very large pectoral-fins that, when rapidly vibrated, give these fish the
ability to "fly." Prochi 1odontids and curimatids are medium to large
sized fishes reminiscent of carps and suckers. They feed on mud, algae
and detritus, and comprise a large percentage of the biomass in most

27
tropical waters. They frequently support productive local commercial
fisheries. Many species make spectacular annual migrations to feeding or
breeding grounds. Anostomids and hemiodontids include mostly herbivores
and pianktivores, with fusiform bodies and conical heads adapted to swim¬
ming in fast currents. The chilodontids are small to medium-sized head-
standers similar to the curimatids, but with teeth.
The differing, still varying taxonomic classifications of characi-
forms exist primarily because current taxonomic groups for the most part
reflect different authors’ attempts to group morphologically similar taxa
together. Differing opinions as to exactly which characters are "similar"
and which are not leads to different conclusions. Although Eigenmann
(1917) expressed much concern about the monophyly of the numerous upper
level taxa he described, the methodologies available at that time were
inadequate. He remarked that the tremendous "radial adaptation" of char-
aciform fishes was at once a paradise for evolutionary biologist and a
nightmare for taxonomists seeking a useful system of names. Durbin Ellis
(1918) came to the conclusion that the genera Hyphessobrycon and Hemigram-
mus were conveniences, and not natural entities (ie. monophyletic units).
Recent efforts, such as those as Vari (1989), to base taxonomy on shared
derived characters and that reflect the phylogenetic history of the fishes
would seem to be the only approach that might eventually lead to agreement
among taxonomists instead of continuing controversy. The characiform
families, and especially the Characidae, contain such divergent groups of
fishes that it is difficult to write a key that is both easy to use and
reasonably precise. The following key should work for most individuals of
most species currently known from the Apure drainage. Readers are also
referred to Taphorn & Lilyestrom (1984) for keys to all the freshwater
families in Venezuela.

28
Key to the Characiform families present in the Apure River drainage.
la. Lips and jaws completely lacking teeth of any kind...
CURIMATIDAE
lb. Teeth present on jaws or lips (teeth on lips may be small and
difficult to see without magnification)... ...2
2a. (lb) Teeth present only on lips, none on jaws ...3
2b. Lips without teeth; upper or both jaws with teeth... ...4
3a. (2a) Gills normal; mouth large and evertible to form a round,
sucking disk; teeth present on edges of lips, teeth small, comb¬
like and numerous (many more than 20 on either side), arranged in
single row at sides of lips, and in two rows near center (midline)
of lips; predorsal spine present, embedded in flesh just anterior
to dorsal fin; large species that exceed 200 mm SL in the first
year of growth... PROCHILODONTIDAE
3b. Gills with fourth arch dilated (expanded), its surface with folds
and wrinkles that mesh with fifth arch; mouth small, not complete¬
ly evertible; teeth larger, not very numerous (less than 20 per
jaw), in single row; no predorsal process present; small species
that seldom exceed 200 mm SL CHIL0D0NTIDAE
4a. (2b) Lower jaw without teeth, or without teeth at center (near
symphysis)... ...5
4b. Lower jaw with complete series of teeth... ...6
5a. (4a) Teeth of upper jaw arranged in semicircle; pectoral fins
not greatly expanded, usually not as long as head; cranium without
fontanels; lower jaw never with teeth at sides, usually completely
lacking teeth... HEMI0D0NTIDAE

29
5b. Teeth of upper jaw arranged in straight line; pectoral fins
greatly expanded, usually longer than head; cranium with large
fontanels; lower jaw usually with a few teeth at sides...
PARODONTIDAE
6a. (4b) Lower jaw trapdoor-like, with extremely large canine teeth,
(the length almost equal to or surpassing eye diameter) that fit
into special "sockets" present in upper jaw and cranium when mouth
is shut... CYNODONTIDAE
6b. Lower jaw without extremely large canines, sometimes with small
canines that measure less than one third of eye diameter, but
usually with multicuspid teeth... ...7
7a. (6b) Adipose fin present... ...8
7b. Adipose fin absent... ...13
8a. (7a) Scales ctenoid; jaws extremely elongate and anal fin with
fewer than fifteen rays... CTENOLUCIIDAE
8b. Scales cycloid; jaws usually not extremely elongate, but if so,
anal fin with more than 15 rays... ...9
9a. (8b) Branquial membranes fused to isthmus for nearly their entire
length; mouth with 6-8 large rabbit-like, sometimes unusually
shaped incisors... ANOSTOMIDAE
9b. Branquial membranes free from isthmus; teeth variable, but not
usually as above... ... 10
10a. (9b) Chest bones (coracoids) greatly expanded to form a deep
narrow keel; dorsal-fin origin behind that of anal fin; lateral
line short, usually directed down towards anal fin origin; pec¬
toral fins greatly enlarged for flight...
GASTEROPELECIDAE

30
10b. Chest usually not greatly expanded, but if so, then dorsal-fin
origin is at a level with, or in front of that of anal fin;
lateral line, if present not usually directed down towards
anal-fin origin; pectoral fins variable... ...11
11a (10b) Lower jaw with two rows of tricuspid teeth (the second row
of small teeth sometimes difficult to see); upper jaw with only
one row of tricuspid teeth and without conical or canine teeth...
LEBIAS INIDAE (Lebiasina)
lib. Teeth not as described above... ...12
12a. Pectoral and pelvic fins enlarged, inserted very low on body and
directed under chest (to support fish resting on substrate);
pectoral fin with the first three or four rays thickened and
unbranched; premaxillary teeth always in a single row; chest and
abdomen flattened (adapted to benthic life as in North American
darters); head conical; mouth small usually subterminal to ventral
(but terminal in a few species)...CHARACIDIIDAE (in part)
12b. (lib) Pectoral and pelvic fins not particularly expanded, usually
inserted near or just below midbody, (pectorals not used to
support body when resting on substrate); first three or four rays
not thickened, usually only first one or two unbranched; premaxil¬
lary teeth in one to three rows, but usually two; chest and abdo¬
men rounded; head not usually conical; mouth variable...
CHARACIDAE (in part)
13a. (7b) Large (up to 1000 mm SL) predators with wide terminal mouths
armed with canine teeth on premaxilla and dentary, and sometimes
on maxilla; scales relatively large (some nearly size of eye)...
ERYTHRINIDAE

31
13b. Small (less than 40 mm SL) fishes with small, superior or
terminal mouths that lack large canine teeth (teeth usually small
to moderate in size, conical or multicuspid); scales relatively
small... ...14
14a. (13b) Dorsal fin origin posterior to that of anal fin; males
with long filamentous extension on opercle...
CHARACIDAE (Corynopoma)
14b. Dorsal fin origin anterior to that of anal fin; males lacking
opercular filament... ...15
15a. (14b) Mouth terminal or subterminal; dorsal fin with fifteen or
more rays; sides with alternating light and dark vertical bars...
CHARACIDIIDAE (Elachocharaxl
15b. Mouth superior; dorsal fin with fewer than fifteen rays; sides
without vertical bars... LEBIASINIDAE (pyrrhulinines)
Anostomidae
The anostomids characteristically are fusiform, somewhat elongated
fishes with conical heads, distinguished from other famil ies by their small,
nonprotractile mouths that usually have eight large incisors arranged in a
single row in either jaw. The anterior nostril is usually tubular, and the
gill membranes are often united to the isthmus (Géry 1977). According to
Winterbottom (1980), this family together with the Chilodontidae, shares
posteriorly replaced, cuspid pharyngeal teeth (not to be confused with the
pharyngeal teeth of Cyprinidae). Roberts (1969), Géry (1977), Winterbottom
(1980) and Vari (1983) have presented very different concepts of subfamil ial
divisions within the Anostomidae. The family comprises about ten genera,
many of which are monotypic. All told, there are at least 100 described

32
species (Géry 1977), of which the largest by far is Leporinus, with some 70
nominal species (Vari 1983).
Most of the remaining taxonomic problems in this family involve the
genus Leporinus, since Winterbottom’s (1980) work has cleared up the
situation in the subfamily Anostominae (comprising the genera Anostomus,
Pseudanos, Gnathodolus, Sartor, and Svnaptolaemus). Neither he nor Vari
(1983) defined for the remaining genera, Abramites Abramoides, Laemolvta.
Leporel1 us, Leporinus, Rhvtiodus and Schizodon, although Winterbottom
(1980) rejected Gery’s (1977) subdivision into two subfamilies, Leporelli-
nae for Leporellus, and Anostominae for all the rest.
The majority of anostomids are medium to large-sized omnivores or
vegetarians that use their incisor-like teeth to clip aquatic vegeta¬
tion, or as forceps to remove prey from crevices. A few anostomids
have developed feeding specializations, these species are relatively
smaller in size, are more elongated and have minute upturned mouths,
often with bizarre dentition.
I have been able to identify twelve species from the Apure drainage,
half of them members of the genus Leporinus. Most of the species’
identifications are still tentative. For those that don’t seem to agree
with any described species, I have used code names.
Very little natural history information is available on anosto¬
mids in the Apure drainage. Most species are probably omnivorous,
though some like Schizodon isognathus tend to be more herbivorous. All
are probably egg scatterers, with no parental care. It is expected
that most of the medium to large species spawn annually at the onset of
the rains. The smaller ones probably spawn repeatedly during the wet
season.

33
Key to the Apure Drainage Species of Anostomidae
This key is based in part on Winterbottom (1980) and Géry (1977),
and will work only for specimens with adult pigmentation patterns.
Since there is considerable size range variation in the family, defining
"juvenile" size for all anostomids is not possible. Juvenile Anostomus
for example, probably obtain adult coloration at a size of 20-30 mm SL,
whereas juvenile Leporinus may not change to the adult pattern until
they reach 100-120 mm. Juveniles all tend to look alike, and are
marked with numerous vertical bars and rounded midlateral spots, that
are gradually lost as the adult pattern is expressed.
la. Caudal fin with oblique black bars; caudal fin densely scaled
for most of its length; anterior nostril not tubular, proximate
in position to the posterior nostril...
Leporellus vittatus (fig. 7)
lb. Caudal fin uniformly pigmented, without bars; caudal fin not
scaled; anterior nostril usually tubular and well separated
from posterior nostril... ...2
2a. (lb) Mouth superior (i.e. upturned) in adults... ...3
(Note: small juvenile Schizodon isognathus have superior mouths
to a size of some 4 cm SL.)
2b. Mouth terminal or inferior in adults... ...5
3a. (2a) Pigmentation pattern comprising a large rounded black spot
surrounded by lighter area located on center of each side, a
smaller spot behind opercle and at base of caudal fin, dorsum
crossed by 8-11 irregular narrow black bars, sides with horizontal
rows of small spots... Pseudanos irinae (fig. 23)

34
3b. Pigmentation pattern not comprising a large rounded spot in center
on the side, but rather of several wide black stripes or one wide
black lateral stripe with several additional horizontal rows of
small dots ...4
4a. (3b) Pigmentation pattern consisting of 10-11 horizontal rows of
small dots on sides and a wide lateral stripe from opercle to tail
(fig. 21) Pseudanos grácil is (fig. 21)
Winterbottom (1980) noted that this species exhibits two color
phases. One is described here, whereas the other (not yet found
in the Apure drainage) consists of 3-4 large rounded spots on mid
sides with horizontal rows of small whitish spots. This is simi¬
lar to the pattern of f\ irinae; however, gracilis never has
horizontal rows of black dots nor dark thin bars crossing the
dorsum.)
4b. Pigmentation pattern consisting of wide black horizontal stripes
separated by narrower white stripes, the widest black stripe
centered on the side and extending from opercle to base of tail...
Anostomus ternetzi (fig. 5)
5a. (2b) Teeth incisiform, with 2-5 cusps, some multicuspid teeth
always present; anal fin usually with only seven branched rays...
Schizodon isognathus (fig. 25)
5b. Teeth incisiform, without cusps or at most bifid, never
multicuspid; anal fin with eight or more branched rays... ...6
6a. (5b) Anal fin usually with ten or more branched rays; body high
and compressed, GBD 37-48% SL; a postventral keel present;
headstanders that usually swim in a vertical position...
Abramites hypselonotus (fig. 3)

35
6b. Anal fin usually with nine branched rays or fewer; body terete, not
very compressed, GBD usually less than 30% SL; no postventral keel;
swimming position horizontal... (genus Leporinus, 6 spp.) 7
7a. (6b) Body with about seven, wide, black vertical bars, the second
usually forming a "Y"; base color yellowish...
Leporinus yophorus (fig. 19)
7b. Body without vertical bars, either plain, with horizontal stripes,
or with large spots; base color variable... ...8
8a. (7b) Body with 4-5 wide black horizontal stripes, separated by
narrower white or yellow stripes...
Leporinus cf striatus (fig. 17)
8b. Body without horizontal stripes, either plain with a small caudal
spot or with large rounded spots on sides... ...9
9a. (8b) Adults with plain brown body and a small black spot at base
of caudal fin... Leporinus sp. (fig. 13)
[lateral-1ine scales 42-44; predorsal scales 12-13; transverse
scales 13-14]
9b. Adults with large black spots on sides of body... ...10
10a. (9b) Adults with five or more large black spots on sides of body,
some along lateral midline, others both above and below these;
mouth inferior... Leporinus cf maculatus (fig. 9)
10b. Adults with only two to four large rounded spots, all along lateral
midline; mouth terminal or subterminal... ...11
11a. (10b) Adults usually with only two large rounded spots along
lateral midline and no faint row of smaller spots on upper sides,
although faint vertical saddles cross dorsum; head and body deeper
and more robust; scales seldom spotted (one spot per scale); body

36
often decidedly higher in front of dorsal fin; eye smaller, in
specimens over 100 mm SL its horizontal diameter less than half of
interorbital width... Leporinus friderici (fig. 11)
[1ateral-1ine scales 38-40; predorsal scales 9-12; transverse
scales 11-12]
lib. Adults usually with three or four large rounded spots along
lateral midline plus a faint row of smaller spots on upper sides;
scales often a spot (one per scale) arranged so as to form hori¬
zontal rows of dots; head and body more slender, not decidedly
higher in front of dorsal fin; eye larger, in specimens over 100
mm SL its horizontal diameter is more than half interorbital
width... Leporinus sp. "aguaro" (fig. 15)
[1ateral-1ine scales about 39; predorsal scales 10-12; transverse
scales 11-12]
Species Accounts
Abramites hypselonotus (Gunther) 1868
High-Backed Headstander - Cabezibajo
Fig. 3. Map: fig. 4. Couplet 6a.
Generic synonymy
Abramites Fowler 1906:331 (type species: Leporinus hypselonotus Giinther
1868, by original designation); Vari & Williams 1987:89 (revision of
genus).
Specific synonymy
Leporinus hypselonotus Giinther 1868a:480 (type locality: Upper Amazon,
Xeberos, Peru); Steindachner 1882:12 (Ciudad Bolivar, Venez.);
Eigenmann 1909:323, 344 (Orinoco); Schultz 1944b:268 (Venez.).

37
Figure 3. Abramites hypselonot.us.

38
Leporinus solarii Holmberg 1887:222 (type locality: Argentina, Rio
Paraná).
Leporinus eaues Boulenger 1896:34 misidentification of hypselonotus
from Brazil).
Abramites hypselonotus Fowler 1906:331 (indicated as type species of
Abramites; 1950:249 (synonymy); Mago L. 1970:75 (Venez.); Géry
1977:175 (key); Vari & Williams 1987:92 (synonymy, key, revision of
genus).
Abramites microcephalus Norman 1926:92 (type locality: near mouth of
Amazon River).
Abramites ternetzi Norman 1926:93 (type locality: Brazil, Matto (= Mato)
Grosso, Sao Luis and Descalvados.
Leporinus salarii Borodin 1929:288, (as a possible synonym of L.
hypselonotus. specific name misspelled).
Leporinus nigripinnis Meinken 1935:193 (type locality: Argentina,
Corrientes).
Abramites eaues Fernández Y. 1950:1 16 (misident ification, Rio
Salinas, Venez.); Mago L. 1970:75 (Venez.).
Abramites solarii Ringuelet, Aramburu and Alonso de Aramburu 1967:213.
Abramites hypselonotus ternetzi Géry 1977:175 (Rio Paraguay Basin).
Abramites hypsel onotus hypsel onotus Géry 1977:75 (Amazon and Orinoco
basins).
Types. (designated by Vari & Williams 1987). Lectotype: BMNH
1867.6.13:40. Paralectotypes: BMNH 1867.6.13:41-42.
Comments. The genus Abramites is comprised of only two species. A.
hypselonotus (Gunther 1868), the only species in Venezuela, has 10-12
branched anal-fin rays and eight, irregularly-shaped bars on the sides

39
between the nape and the caudal peduncle. A. eques (Steindachner) 1878,
has 13-14 branched anal-fin rays and only five bars on the body, the
anteriormost situated under the dorsal fin (Vari & Williams 1987).
Etymology. ABRAM = from Abramis, a genus of similarly shaped fish,
ITES = like; HYPSELO = raised or high, NOTUS = back.
Description
Illustrations. Fig. 3; Axelrod et al. 1971:F-2.10; Vari & Williams
1987:figs. 2-5.
Diagnosis. The unusual body shape and pigmentation pattern (fig. 3)
are sufficient to identify this species. It is further distinguished by
a terminal mouth, a deep, compressed body (GBD 37-48% of SL), and the
presence of a postventral keel.
Size. It grows to about 130 mm SL.
Morphology. See Vari & Williams (1987). The conical head abruptly
expands vertically behind the eye and the dorsal profile rises steeply
from the nape to the dorsal-fin origin to give this species its charac¬
teristic high-backed appearance. The body is more compressed than in
most anostomids.
Counts. DR 10-13 (up to three unbranched); AR 12-15 (up to three
unbranched); PR 13-15; VR 8-9, 37-38 vertebrae.
Pigmentation. The body is tan or silvery with eight irregular
vertical bars, the darkest and widest bar extends from the pelvic fins
across body and onto the anterior dorsal fin. There is also a black
stripe from the tip of the lower jaw through the eye, which continues
obliquely upward toward the dorsum. The adipose fin is yellow and
black, the caudal and pectoral fins are clear, and the anal fin is
bl ack.

Figure 4. Apure drainage distribution of Abramites hypselonotus.

(«cal* •■•rica

42
Distribution and Natural History
Range. A. hypselonotus occurs in the Amazon, Orinoco, Paraguay and
lower Paraná basins (Vari & Williams 1987).
Apure distribution. Map: fig. 4. Although few specimens were
collected, the localities range across the entire Apure drainage.
Habitat. This species was most frequently collected in habitats
with aquatic vegetation in quiet sidewaters of large Whitewater rivers
in the low llanos. It swims in a head down position, often aligning
itself with the stems of plants.
Abundance. UNCOMMON.
Number of specimens examined. 28 from 14 collections.
Food. OMNIVORE. Its diet includes aquatic invertebrates, as well
as vegetable matter.
Reproduction. Probable strategy: rl.
Importance. This is a highly valued ornamental.
Anostomus ternetzi Fernández Y. 1949
Red-Mouthed Headstander - Cabezibajo Bocaroja
Fig. 5. Map fig. 6. Couplet 4b.
Generic synonymy
Anostomus Gronow 1763:122 (description and figure, no species mentioned,
work not accepted by ICZN).
Anostomus Scopol i 1777:451 (no species mentioned, but since this work
was based on Gronow the type species is that illustrated by Gronow
1756, which is Salmo anostomus Linnaeus).
Anostoma Rafinesque 1815 (alternative spelling).
Mormvrhvnchus Swainson 1839:186, 291 (type species: Mormvrhvnchus
gronovii Swainson = Salmo anostomus Linnaeus, by monotypy).

43
Figure 5. Anostomus ternetzi.

44
Histriodromus Gistel 1848:8 (type species: Salmo anostomus Linnaeus,
by monotypy).
Pithecocharax Fowler 1906:319 (type species: Salmo anostomus Linnaeus,
by original designation, this name was proposed as a substitute for
Anostomus Walbaum 1792).
Specific synonymy
Anostomus ternetzi Fernández Y. 1949:293 (type locality: Orinoco River
System); Kndppel 1972:231 (diet).
Anostomus anostomus (non Linnaeus); Myers 1950:184 (part).
Types. Holotype: Museo de Ciencias Naturales de Caracas 46001.
Paratypes: MCNC 46002; CAS 20093(1), 20094(1), 20095(4), 20096(1).
Etymology. ANO = upturned, ST0MUS = mouth; TERNETZI = for Carl
Ternetz.
Description
Illustrations. Fig. 5; Axelrod et al. 1971:F-40.11 - 40.12; Géry
1977:188; Winterbottom 1980:81 fig. 17.
Diagnosis. A. ternetzi is separable from Leporinus by the superior
mouth. The color pattern is distinctive for the species. The white
stripe that extends from between nostrils along the dorsal midline to
the dorsal-fin origin in this species, is dark in A^_ anostomus. a spe¬
cies that occurs in other parts of Venezuela.
Size. It grows to about 100 mm SL.
Counts. This species has three branchiostegal rays instead of the
four usually found in other Anostomus species. DR 12-13; AR iii7-iii8;
PR i13-i15; VR i8; LS 39-42; PDS 11-14; CPS 16.
Pigmentation. The basic pigmentation pattern consists of three wide
black horizontal stripes separated by narrower white zigzag stripes.

45
The widest black stripe is situated at midside and extends from the
opercle to the base of the tail. Another starts at the pectoral-fin
insertion and extends back to the lower part of the caudal peduncle.
There is a narrow black stripe situated on either side of the narrow
white stripe that runs along the ventral midline, and another wide black
stripe on either side of the white stripe that extends from between the
nostrils to the dorsal fin origin. In many individuals the black (or
dark brown) pigment uniformly covers most of the dorsum, although it is
sometimes divided into several rows of dots by lighter areas. Fins that
are normally reddish in life, would appear clear in preserved specimens.
Distribution and Natural Hi story
Range. This species occurs in the Amazon and Orinoco basins, and in
Guyana.
Apure distribution. Map: fig. 6. It is known only from northern
Apure state, and the Aguaro-Guariquito system.
Habitat. It is found in black or Clearwater streams with abundant
aquatic vegetation, in the sandy-soiled areas of northern Apure state
and western Guárico.
Abundance. UNCOMMON.
Number of specimens examined. 69 from 19 collections.
Food. OMNIVORE. This fish probably feeds on vegetable material as
well as aquatic insects. It swims head down, and presumably feeds in
this position amidst aquatic vegetation.
Reproduction. Probable strategy: rl. Winterbottom (1980) noticed
some sexual dimorphism, in that males have a greater caudal peduncle
depth. Sterba (1972) stated that the similar A. anostomus is territori¬
al. Axelrod et al. (1971) stated that in A. anostomus the pair quivered

Figure 6. Apure drainage distribution of Anostomus ternetzi.

CUENCA OEL RIO APURE
Ct CAI
• AAflCl
too
-p*
•^1

48
side by side, and expelled a large number of eggs all over the glass,
plants and the bottom of the aquarium. Fry were slender, about 15 mm
long, and swam in a normal horizontal position.
Importance. Ornamental.
Leporellus vittatus (Valenciennes) 1849
Stripe Tailed Leporellus - Mije de Cola Rayada
Fig. 7. Map fig. 8. Couplet la.
Generic svnonvmv
Leporel1 us Lütken 1874b:129 (type species: Leporinus pictus Kner, by
monotypy).
Leporinodus Eigenmann 1922:116 (type species: Leporinodus retropinnus
Eigenmann 1922, by original designation, (described in footnote).
Specific synonymy
Leporinus vittatus Valenciennes jn Cuvier & Valenciennes 1849:59 (type
locality: Rio Amazonas).
Leporel1 us vittatus Lütken 1874b:129; Fowler 1950:253 (synonymy);
Géry 1977:151 (diagnosis).
Leporinodus vittatus Eigenmann 1922:117.
Leporinus pictus Kner 1859:172, pi. 8, fig. 19 (type locality: Orissanga,
Est. de Sao Paulo).
Salmo cagoara Natterer in Kner 1859:172 (name in synonymy).
Leporinus maculifrons Reinhardt jn Lütken 1874b:201 (name in synonymy).
Leporellus timbore Eigenmann 1922:117 (type locality: Rio das Velhas).
Comments. Géry (1977) listed seven forms in this genus, of which
he listed (without explanation) L. maculifrons. L. timbore, and L.
cartledgei as synonyms of L. pictus. None are well known, and a revi¬
sion of the genus is necessary before any further comments can be made

49
Figure 7. Leporellus vittatus.

50
about synonymies. I have used the synonymy of Fowler (1950) here, but
additional study may change these.
Etvmology. LEPORELLUS = like a rabbit, referring to the teeth;
VITTATUS = spotted.
Description
Illustrations. Fig. 7; Géry 1977:153.
Diagnosis. This is the only anostomid with a scaled, obliquely
barred caudal fin. The pigmentation pattern of the body (fig. 7) is
also distinctive.
Size. It grows to at least 200 mm SL.
Morphology. It is elongate with a large head and eye. The mouth is
subterminal with the lower lip reverted to form a wide fold. The gill
membranes are joined to the isthmus. The nostrils are proximate, and
not tubular (unusual in this family). The caudal fin is completely
scaled.
Counts. DR 13; AR 10-11; PR 16-17; VR i8; LS 42; PDS 12-13; CPS
16-17; TS 11-12.
Pigmentation. The base color of the body is yellow-brown in life.
The white tail has a black stripe through the central rays, and two
black oblique bars in each lobe. The top of the head, dorsum, and upper
sides are heavily spotted, but the belly is white. The spots on the
sides (one per scale) are arranged in nine or ten horizontal rows. The
sides have two white stripes devoid of spots that continue posteriorly
onto the tail. The dorsal fin is white with a large black distal blotch
on its first rays, and a row of black spots nearer base. The pectoral
and pelvic fins are whitish to clear. The anal fin is white with a
central black blotch.

Figure 8. Apure drainage distribution of Leporellus vittatus.

en
ro

53
Distribution and Natural Hi story
Range. The range is not well documented but it probably occurs in
most of the Andes in Venezuela and Colombia. There are reports of this
species from the Guyana Shield in Venezuela, but I have not seen speci¬
mens. The form there may represent a different species.
Apure distribution. Map: fig. 8. It has been found only in the
western portion of the Apure drainage, usually in the Andean foothills.
Habitat. It lives in clear, clean, fast-flowing mountain streams.
Juveniles have been taken at Bruzual in the quiet edges of the Apure
River from aquatic vegetation. They perhaps were washed out of the
mountains, or the reproductive cycle might include a downstream phase
with later upstream migration.
Abundance. UNCOMMON.
Number of specimens examined. 49 from 16 collections.
Food. OMNIVORE. It feeds on both insects and vegetable matter. I
observed several individuals biting at rocks in the clear waters of the
upper Rio Caparo, presumably selecting items from the epibenthos.
Reproduction. Probable strategy: r2.
Importance. It has great potential as an ornamental.
Leporinus cf maculatus Müller & Troschel 1844
Spotted Leporinus - Mije Pintado or Cabeza de Manteco
Fig. 9. Map: fig. 10. Couplet 10a.
Generic synonymy
Leporinus Spix i_n Agassiz 1829:65 (type species: Leporinus fasciata
Agassiz, by monotypy).
Hypomasticus Borodin 1929:287 (type species: Leporinus mormvrops
Steindachner 1875, designated by Fowler 1950:228).


55
Specific synonymy
Leporinus maculatus Müller & Troschel 1844:86 (type locality: Guyana);
1845:11 (rivers of Guyana); Eigenmann 1912a:305 (redescription);
Fowler 1950:237 (synonymy); Géry 1977:166 (key, discussion of many-
spotted Leporinus).
Leporinus megalepis Gunther 1863:443 (type locality: Essequibo River,
Guyana).
Leporinus marcqravii Lütken 1874b:130 (type locality: Rio Velhas and
tributaries).
Comments. The counts for Apure drainage specimens are consistently
higher than those given by Eigenmann (1912) for specimens from Guyana,
either because of a more inclusive counting method, or because different
species are involved. Géry (1977) discussed the complex taxonomic
problems of the many-spotted Leporinus species. He stated that in
current usage both aquarists and ichthyologists have confused L. pel 1e-
grini with L. maculatus, and that the name L. maculatus has been applied
to at least two very different species. I follow him in using L. macu-
latus for the species with a ventral mouth. He placed this species in
the subgenus Hypomasticus.
Etymology. LEPO = rabbit, RINUS = snout; MACULATUS = spotted.
Description
Illustrations. Fig. 9; Eigenmann 1912:pi. 43, fig. 2; Axelrod et
al. 1971:F377.00; Géry 1977:166; Román 1985:141.
Diagnosis. The pigmentation pattern is distinctive (fig. 9).
This is the only Leporinus with five or more large spots on the sides.
The ventral mouth is also unique.
Size. It grows to at least 200 mm SL.

56
Morphology. The head length is about equal to the greatest body
depth. The interorbital width is less than the snout length.
Counts. DR i i 10; AR Ü9; PR i 14-i 16; VR i8; CPS 16; LS 38-40; TS
11-12 5-6 above, and 5 below lateral line.
Measurements. HL = 22% SL; GBD = 27-33% SL (Eigenmann 1912 for
Guyanan specimens). In Apure specimens, GBD and HL = 24-26% SL; EYE =
50-66% HL, 10 = 43-55% HL, and SNT = 20-24% HL.
Pigmentation. The body is marked with several large ovate black
spots along the lateral midline, and on the upper and lower sides This
is a pattern similar to that of most juvenile anostomids, but in this
case it has been retained in the adult.
Distribution and Natural Hi story
Range. It occurs in the Guianas and in the Orinoco Basin.
Apure distribution. Map: fig. 10. It is known only from the Agua-
ro-Guariquito system in the easternmost portion of the Apure drainage.
Habitat. It occurs in clear and blackwater creeks of the upper
Aguaro River system, in sandy bottomed savanna streams with abundant
aquatic vegetation.
Abundance. RARE.
Number of specimens examined. 6 from 4 collections.
Food. HERBIVORE. It is probably mainly herbivorous, though occa¬
sionally aquatic insects may be included in the diet.
Reproduction. Probable strategy: r2. It is probably an egg scatterer
that synchronizes its annual reproductive effort with the beginning of the
rainy season. Axelrod et al. (1971) reported that this species spawned in
aquaria, scattering eggs all over, and then eating them if not removed.
Importance. Ornamental.

Figure 10. Apure drainage distribution of Leporinus cf maculatus.

10“
9°
LA UNIVERSIDAD O'if SiEMfif '
UNIVERSIDAD NACIONAL EXPERIMENTAL
DE LOS LLANOS OCCIDENTALES
"EZ fcQUIEL ZAMORA"
VICE RECTORADO EN PORTUGUESA
72°
CUENCA DEL RIO APURE
en
00

59
Leporinus friderici (Bloch) 1794
Leporinus - Mije
Fig. 11. Map: fig. 12. Couplet 11a.
Specific synonymy
Salmo friderici Bloch 1794:94, pi. 378 (type locality: Surinam).
Leporinus friderici Müller & Troschel 1844:87; 1845:11; Eigenmann &
Allen 1942:305 (Venezuela); Fowler:1950:233 (synonymy).
Leporinus fridericii Kner 1859:170 (misspelling).
Leporinus frederici Valenciennes i_n Cuvier & Valenciennes 1849:25
(misspelling).
Leporinus fradericii Goeldi 1898:482 (misspelling).
Curimatus frederici Perugia 1891:641.
Curimatus acutidens Valenciennes 1847:9, pi. 8, figs 1-la (type locali¬
ty: South America)
Leporinus megalepis (non Günther 1863) Günther 1864:307 (part).
Leporinus 1 eschenaultii Valenciennes i_n Cuvier & Valenciennes 1849:30,
pi. 635 (type locality: Mana); Peters 1877:472 (Calabozo, Venezuela);
Eigenmann & Eigenmann 1891:51 (Calabozo, Venezuela); Pellegrin
1899:157 (Apure River, Venezuela).
Comments. Identification of species in the L. friderici group is
difficult, see Géry (1977) and Garavello (1988) for a brief outline of
the problem. Apure specimens resemble L. subniqer Fowler 1943 from
Florencia, Colombia in the Andean piedmont, and both of these seem
quite similar to L. bahiensis Steindachner 1875 from Brazil.
Etvmoloqy. FRIDERICI = probably named for the German monarch
Frederick the Great (1712-1786).

60
Figure 11. Leporinus friderici.

61
Description
Illustrations. Fig. 11; Eigenmann 1912:pi. 63 fig. 4; Axelrod et
al. 1971:F-376.00; Géry 1977:168.
Diagnosis. The pigmentation pattern is distinctive (fig. 11). Adults
usually have only two large spots on the sides. This species is very simi¬
lar to Leporinus sp. "aguaro," but that form usually has three large spots
on the sides, and has a relatively larger eye (the eye diameter is more than
half of the interorbital width vs less than half in U. friderici. Faded
specimens of L_ friderici (without spots) are similar to "Leporinus sp.
but the latter has much wider lips, and more lateral-line scales (42-44 vs
38-40). L. boehlkei Garavello 1988 has fewer lateral-line scales (35-36),
and fewer circumpeduncular scales (12 vs 16).
Size. It reaches at least 350 mm SL.
Morphology. This is a very robust, deep-bodied Leporinus with a
stocky appearance. The mouth is terminal.
Counts. DR i i 10; AR Ü9; PR i 14-i 16; VR i8; LS 38-40; PDS 9-12;
CPS 16; TS 11-12; teeth 3-3 on premaxilla, middle incisors bifid, the
others strangely curved; 3-4 teeth on dentary, the last one tiny and
next to base of last big tooth.
Measurements. GBD 32-36% SL.
Pigmentation. The dorsum is grayish, and crossed by about 12-15
faint vertical bars and the ventrum is light tan. The sides are brown,
with at least two large, rounded black spots, the first spot under
dorsal fin base, the second just anterior to the adipose fin. Sometimes
there is an additional spot at the base of the caudal fin. The fins,
especially the anal, are dusky and heavily pigmented on the membranes.
The adipose fin is often edged in black.

Figure 12. Apure drainage distribution of Leporinus friderici.

CUENCA DEL RIO APURE
O â– 
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64
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins, and the
Guianas.
Apure distribution. Map: fig. 12. It is common throughout the
piedmont and llanos.
Habitat. It lives in a wide variety of streams and rivers from
both white and blackwater habitats.
Abundance. COMMON.
Number of specimens examined. 230 from 64 collections.
Food. OMNIVORE. It feeds mostly on aquatic plants, but seeds and
carrion (fish flesh) are also included in its diet (K. Winemiller pers.
com.).
Reproduction. Probable strategy: r2. This species migrates down¬
stream shortly after the first rains of the season. Spawning probably
takes place in midriver, in a fashion similar to Prochi 1odus mariae.
Migrations. This fish is known to accompany Prochilodus mariae on
annual migrations from the lowland floodplain habitats upstream to
piedmont and montane streams at the beginning of the dry season, and
thence back downstream when the seasonal rains begin.
Importance. It is of minor importance as a commercial species.
Leporinus sp.
Fatlip Leporinus - Mije
Fig. 13. Map fig. 14. Couplet 9a.
Comments. This may be a new species. Its plain coloration and
meristic characters don’t agree well with any described species.

65
Figure 13. Leporinus sp.

66
Description
111ustrations. Fig. 13.
Diagnosis. This fish is quite plain, usually brown when adult,
with a small black spot at the base of the tail (fig. 13). The lip is
much thicker and more gently curved than in the similar L. friderici.
It has a more slender body as well. The lateral-line scale count of 42
to 44 is higher than for most other Apure drainage Leporinus species, in
which the number usually does not exceed 40.
Size. It reaches at least 250 mm SL.
Counts. DR ii10; AR ii9; PR i14-i18; VR i8; LS 42-44; PDS 12-13;
CPS 16; TS 13-14.
Measurements. GBD 26-30% SL in specimens above 90 mm SL.
Pigmentation. The body in adults is plain brown with a small spot
at the base of the caudal fin. Smaller specimens may have two or three
spots on the sides.
Pistribution and Natural Hi story
Range. It is known from the Orinoco Basin.
Apure distribution. Map: fig. 14. It occurs throughout the pied¬
mont and llanos.
Habitat. It lives in muddy (whitewater) streams and rivers.
Abundance. COMMON.
Number of specimens examined. 81 from 25 collections.
Food. OMNIVORE. It is probably mostly vegetarian, but undoubt¬
edly includes some aquatic invertebrates in its diet.
Reproduction. Probable strategy: r2.
Migrations. It makes annual migration similar to those of
Prochi 1odus mariae, and often accompanies that species during the

Figure 14. Apure drainage distribution of Leporinus sp.

CUENCA DEL RIO APURE
<00
cr>
oo

69
"ribazón" (the spectacular annual upstream migration that begins with
the onset of the dry season in October or November).
Importance. It is of only minor commercial importance.
Leporinus sp. "aguaro"
Leporinus - Mije
Fig. 15. Map fig. 16. Couplet lib.
Comments. This species is similar to the species I have identified
as L. friderici, and could turn out to be the "real" L. friderici. If
that is the case, the species I have identified as L. friderici is
probably new. The original descriptions don’t provide adequate informa¬
tion to make a certain determination. L. friderici was described from
Surinam, presumably from blackwater rivers typical of the Guyana Shield
and coastal drainages. Leporinus sp. "aguaro," is apparently restricted
to similar habitats in the Apure drainage.
Description
Illustrations. Fig. 15.
Di agnosis. This species can be distinguished by pigmentation
pattern (fig. 15), which consists of three to four large spots along the
lateral midline plus a faint row of smaller spots above them on the
upper sides. It has fewer lateral scales (39 vs 42-44) than Leporinus
sp., and a relatively larger eye (the eye diameter greater than half of
the interorbital width) than L. friderici (less than half the interor¬
bital width).
Size. It grows to at least 200 mm SL.
Morpholoov. The snout length is about equal to the interorbital
width.

70
Figure 15. Leporinus sp. "aguaro".

71
Counts. DR ii10; AR Ü9; PR i15-i16; VR i8; LS 37-39; PDS 10-12;
CPS 16; TS 11-12, with 5-6 scales above and five below the lateral
line.
Measurements. GBD 29-32% SL; HL 28-31% SL; EYE in HL 22-34%, in
SNT 53-96%, in 10 54-86% (the large variation in eye diameter is due to
allometry, juveniles have relatively larger eyes).
Pigmentation. The body is brown, darkest on the dorsum and
becomes gradually whiter towards the belly. The sides are marked with
three or four large black rounded spots, the first under the posterior
dorsal-fin base, the second just anterior to the adipose fin, the third
under the adipose fin, and the fourth on the caudal peduncle. Another
series of seven smaller spots is situated above these on the upper
sides. Each scale has a dark spot on its margin, resulting in up to 10
horizontal rows of dots; in some specimens the dark pigment completely
outlines each scale, forming a net-like pattern.
Pistribution and Natural Hi story
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 16. It is known only from the
Aguaro River system.
Habitat. It occurs in blackwater streams and rivers.
Abundance. RARE.
Number of specimens examined. 3 from 3 collections.
Food. OMNIVORE. It is probably mostly herbivorous.
Reproduction. Probable strategy: r2.
Migrations. They are probable, but not documented.
Importance. It is potentially valuable as an ornamental.

Figure 16. Apure drainage distribution of Leporiñus sp. "aguaro

LA uNIVtASIOAO Q'lí SltMBÍl
UNIVERSIDAD NACIONAL EXPERIMENTAL
DE LOS LLANOS OCCIDENTALES
"EZEQUIEL ZAMORA”
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CUENCA DEL RIO APURE
MC4LA •■«re»
^4
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Leporinus cf striatus Kner 1859
Striped Leporinus - Mije Rayado
Fig. 17. Map fig. 18. Couplet 8a.
74
Specific Synonymy
Leporinus striatus Kner 1859:171 pi. 8, fig. 18 (type locality:
Orissanga, Río Paraná and Caicara Mato Grosso, Brazil); Pellegrin
1899:157 (Apure River, Venezuela); Géry 1977:170 (key).
Ledorinus (error) striatus Bertoni 1939:54 (Paraguay).
Salmo tiririca Natterer i_n Kner 1859:172 (?based on same type as
L. striatus Kner).
Comments. Since L. striatus was described from southern Brazil, it is
doubtful that the same species is present in the Apure drainage. Fowler’s
illustration (1950) depicts a fish with only two stripes on the side. I
have examined the holotype (FMNH 53366) of the similar L. arcus Eigenmann
1912, which was described from British Guiana, and which would perhaps be
more 1 ikely to occur in Venezuela. It is, however, a different-looking fish
with only four stripes on either side (vs five in Apure specimens). In
addition the dorsum in the L. arcus holotype is uniformly dark, and though
the pigment may have obscured a stripe that may have originally been
present, none is shown in the figure of the holotype (Eigenmann 1912:pi. 62,
fig. 3). The holotype has 36 lateral-line scales (vs 38 in Apure specimens)
and ten transverse scales (vs 12). Finally, specimens of the type series of
L. arcus greatly exceed the maximum recorded size for L. striatus. These
data strongly suggest that the Apure specimens are not L. arcus. I suspect
that the Apure drainage form is undescribed, but this cannot be finally
resolved until it can be compared with fresh material from the type locali¬
ties of L. arcus and L. striatus.

75
Figure 17. Leporinus cf striatus

76
Etymology. STRIATUS = striped.
Description
Illustrations. Fig. 17; Axelrod et al. 1971:F-378.00; Román
1983:99.
Di agnosis. The color pattern of black stripes over a yellowish
background (fig. 17) is unique for Apure drainage anostomids. In this
regard it is superficially similar to Anostomus ternetzi, but that
species has a superior mouth and is much more slender and elongate.
Size. It is seldom found over 100 mm SL, but can reach 175 mm.
Morphology. This is a robust, stocky fish. The horizontal diame¬
ter of the orbit is longer than the snout length, but slightly less than
the interorbital width. The mouth is subterminal.
Counts. DR i i 10; AR Ü8; PR i 14; VR i8; LS 38; PDS 10; CPS 16; TS 12;
teeth 3-3, 3-3 or 4-4 (if four the last very small), the teeth are curved
incisors with brown tips, not truly cuspid but with irregular margins. The
center of the roof of the mouth has a bulbous papillose projection.
Measurements. GBD about 29% SL; HL 36% SL; EYE 36% HL. IO 39% HL.
Pigmentation. Juveniles up to about 40 mm SL show vertical
banding instead of the typical adult pattern of horizontal light and
dark stripes. They also have a jet-black adipose fin with a light
center, a dark crescent along base of tail with darker dot at midpoint,
and a black anal fin. Juveniles also lack pigment on the fontanel and
have about three vertical stripes anterior to the dorsal fin. At 19 mm
SL one juvenile had only vertical barring, but another specimen of 37
mm SL had the beginnings of horizontal stripes.
Adults have a black line along the dorsal midline of the body in
front of the dorsal fin, but the dorsal midline of the head has a white

77
stripe. They have five additional dark stripes on either side, the
first of which extends from the occiput along the uppermost sides and
back to the adipose fin. The second extends from the snout, back above
the eye to the uppermost caudal-fin base. A third extends from the
snout through the eye across the opercle and back along the midside to
midtail. The fourth extends from the pectoral-fin base (which is dark
dorsally near its base) back to the anal fin and onto the lower caudal
peduncle. The fifth stripe is faint and extends from the lower pectoral -fin
base to the pelvic-fin base. The adipose and anal fins are black, the other
fins white, tan, or transparent.
Distribution and Natural Hi story
Underwater observations confirm that this species usually travels
in schools, as do most Leporinus. However, since they occur over rocky
substrates where seines are easily avoided, most of our collections are
of very few specimens.
Range. It has been reported from the Orinoco, Amazon and Paraná
basins.
Apure distribution. Map: fig. 18. It is most common in the Andean
piedmont and streams in the upper llanos.
Habitat. U. striatus is most commonly found in clear streams and
rivers, with fast currents and rocky substrates.
Abundance. COMMON.
Number of specimens examined. 109 from 35 collections.
Food. OMNIVORE. It is probably mostly herbivorous.
Reproduction. Probable strategy: r2.
Importance. Ornamental.

Figure 18. Apure drainage distribution of Leporinus striatus.

I'
10"
9»
8*
CUENCA DEL RIO APURE
IOO
vo

80
Leporinus vophorus Eigenmann 1922
Banded Leporinus - Mije Rayada
Fig. 19. Map fig. 20. Couplet 8a.
Specific synonymy.
Leporinus yophorus Eigenmann 1922:233 (type locality: Barrigón,
Colombia, upper Rio Meta)
?Leporinus affinis Pellegrin 1899:157 (Apure River, Venez.).
Types: Holotype: CAS 61680 (formerly III 15025).
Comments. There are many names available for vertically banded Lepori -
nus species. Most often they are called L. fasciatus, but L. 1 atofasciatus
and L. pearsoni are other possibilities for Venezuelan specimens. The Apure
form agrees almost exactly with Eigenmann’s (1922) original description of
L. yophorus. Since both the habitat and drainage coincide, I feel reasona¬
bly confident about this identification. It is also possible that L. yopho¬
rus could prove to be a junior synonym of one of the older names such as
Leporinus affinis Giinther 1864, which was described from Para, Brazil, and
was cited by Pellegrin (1899) from the Apure River. However, that species,
which has nine vertical bars according to Géry (1977), is probably not
present in the Orinoco Basin.
Etymology. Y = the letter "Y" refers to the Y-shaped bar on the
side, 0PH0RUS = bearer of.
Description
Illustrations. Fig. 19; Eigenmann 1922:317 pi. 20, fig 4.
Diagnosis. The color pattern (fig. 19) is distinctive since this
is the only Apure drainage Leporinus with bold vertical bars.
Size. It grows to about 180 mm SL

81
Figure 19. Leporinus vophorus.

82
Morphology. This is an elongate, slender Leporinus with a terminal
mouth.
Counts. DR ii10; AR Ü9; PR i 15-i 16; VR i9; LS 42-45; PDS 14-16;
CPS 16; TS 12, 6 above, 5 below lateral line.
Measurements. Eigenmann (1922) gave the following measurements for
the type: Head 5.0 in SL, depth 4.66, eye 2 in snout, 4.5 in the head,
2.0 in the interorbital; isthmus 3.5 in head; depth of caudal peduncle
2.4 in head. Apure specimens: GBD 23% (adults) to 29% (juveniles) SL;
Eye 18% (adults) to 29% (juveniles) HL.
Pigmentation. The body is yellowish with seven wide vertical bars.
The second bar is divided at the level of the lateral line to form a "Y"
that extends dorsally.
Distribution and Natural History
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 20. It is known only from four
localities, three from within or near the Apure River itself, and one
from the Tucupido River in the piedmont.
Habitat. There are insufficient numbers of collections to
permit a thorough characterization of this species’ habitat.
The one adult captured was collected from a site in the Andean piedmont
in a rapidly flowing, rocky river. The juvenile specimens were taken
from near shore in muddy-bottomed side pools of the Apure River near
Bruzual, amidst aquatic vegetation and submerged grasses. Thus, it
seems that llaneran whitewater rivers are this species habitat.
Abundance. UNCOMMON, but it is probably difficult to capture.
Number of specimens examined. 84 from 11 collections.
Food. OMNIVORE. It is probably mostly herbivorous.

Figure 20. Apure drainage distribution of Leporinus yophorus.

UNIVERSIDAD NACIONAL EXPERIMENTAL
DE LOS LLANOS OCCIDENTALES
"EZEOUIEL ZAMORA'
CUENCA OEL RIO APURE
licait ••»*«:»
í 00
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85
Reproduction. Probable strategy: r2.
Importance. Ornamental.
Pseudanos gracilis (Kner) 1859
Banded Headstander - Cabezibajo Rayada
Fig. 21. Map fig. 22. Couplet 4b.
Generic synonymy
Pseudanos Winterbottom 1980:24 (type species: Schizodon trimaculatus
Kner 1859:161, by original designation).
Specific synonymy
Schizodon grácil is Kner 1859:160 (type locality: Rio Guaporé, Brazil).
Anostomus grácil is Myers 1950:184; Knoppel 1972:268 (diet); Mago L.
1970:75.
Pseudanos grácil is Winterbottom 1980:24 (redescription).
Types. Holotype: NMW 57-119.
Description
Illustrations. Fig. 21; Winterbottom 1980:83 fig. 20.
Diagnosis. The minute, superior mouth distinguishes this species
from all Leporinus species. The pigmentation pattern (fig. 21) is also
characteristic, and useful to distinguish it from P. grácilis, and
Anostomus ternetzi.
Size. It grows to 160 mm SL, but most individuals are smaller.
Morphology. The body is fusiform but somewhat compressed.
Counts. DR i ii10; AR iii8; PR i13-i16; VR i8; LS 43-47; PDS 11-15;
CPS 16; TS 10, teeth 4-4 in both jaws (each with 2 to 4 cusps, though
these can wear down, and become less conspicuous).
Measurements. GBD 18.3-25.4% SL

86
Figure 21. Pseudanos grácil is.

Figure 22. Apure drainage distribution of Pseudanos gracilis.


89
Pigmentation. The pigmentation pattern varies in preserved material,
but the dorsum never has cross bars. The body has up to 11 thin horizontal
stripes formed by rows of dots (one per scale). A median predorsal stripe
is also present. The midlateral sides are marked with a dark stripe or with
series of up to four large rounded spots.
Etymology. PSEUD = false, ANOS = the first four letters of Anostomus.
a very similar genus; GRACILIS = slender.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map: fig. 22. It has been found only in the
Aguaro River system. Interestingly, the distribution of this species seems
to complement that of P. vrinae, which is absent from the Aguaro systems but
present in suitable blackwater habitats of northern Apure and Barinas.
Habitat. It is restricted to blackwater streams with abundant aquatic
vegetation.
Abundance. RARE.
Number of specimens examined. 8 from 5 collections.
Food. OMNIVORE: It feeds on plant remains, fungi, algae, detritus,
sand and terrestrial insects (Winterbottom 1980).
Reproduction. Probable strategy: rl.
Importance. Ornamental.
Pseudanos irinae Winterbottom 1980
Three spotted Headstander - Cabezibajo de Tres Puntos
Fig. 23. Map fig. 24. Couplet 3a.
Specific synonymy
Pseudanos irinae Winterbottom 1980:27 (type locality: Orinoco River
system, Caño de Quiribana where it empties into the Orinoco).

90
Figure 23. Pseudanos irinae.

91
Anostomus trimaculatus (non Kner 1859) Eigenmann 1912:295 (Essequibo
River; Myers 1950:184 (upper Orinoco); Ramirez 1957:157 (Venez.);
Mago L. 1970:75 (Venez.).
Types. Holotype: CAS 58809. Paratypes: deposited at AMNH, BMNH,
CAS, FMNH (3) (Winterbottom 1980), catalog numbers not given.
Etvmoloqy. IRINAE = named for the wife of Dr. R. Winterbottom,
Irina.
Description
Illustrations. Fig. 23; Axelrod et al. 1971:F-41.00 (identified as
Anostomus trimaculatus); Winterbottom 1980:83, fig. 21.
Diagnosis. The pigmentation pattern is distinctive (fig. 23).
Size. It grows to about 100 mm SL.
Morphology. The body is fusiform and somewhat compressed.
Counts. DR 13-14; AR iii7-iii8; PR i 13-i16; VR i7-i8; LS 41-45;
CPS 16.
Measurements. GBD 23-29% SL.
Pigmentation. This fish is yellow-brown, paler below, with the dorsum
crossed with irregular, thin vertical bars, and two to four dark spots or
blotches on the sides. The first of these spots covers scales three to five
of the 1 ateral- line series and is situated partly beneath the 1 ateral line),
the other spots cover scales 16-19, 29-30, and 41-46. The first and third
spots are often faint and may be absent. The area around each spot is
1ighter.
Pistribution and Natural History
Range. It occurs in the Orinoco and Essequibo basins, of Colombia,
Venezuela and Guyana.

Figure 24. Apure drainage distribution of Pseudanos irinae.

CUENCA DEL RIO APURE
Ola*
(«CALA «MAflCA
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94
Apure distribution. Map: fig. 24. It is so far known only from the
Suripá River in Barinas, and the Guaritico system of northern Apure state.
Habitat. It is restricted to black or Clearwater streams in the
savannas of the low llanos, and is usually found in aquatic vegetation.
Abundance. UNCOMMON.
Number of specimens examined. 17 from 6 collections.
Food. OMNIVORE. The diet of this species is probably simi 1 ar to that
of the similar, preceding species which feeds on plant remains, fungi,
algae, detritus, and insects.
Reproduction. Probable strategy: rl.
Importance. Ornamental.
Schizodon isognathus Kner 1859
Boquimí, Coti, Pijotero - Schizodon
Fig. 25. Map: fig. 26. Couplet 5a.
Generic synonymy
Schizodon Agassiz 1829:66 (type species: Curimatus fasciatus Agassiz, by
monotypy).
Specific synonymy
Schizodon isognathus Kner 1859:163 pi. 6 (not pi. 7 as stated in
the text; type locality: Rio Cujaba).
Anostomus isognathus Peters 1877:472 (San Fernando de Apure, Venez.).
Comments. Recent investigations by biologist Maria Esther Antonio
of the Universidad Central de Caracas indicate that this species is
distinct from the Amazonian S_l. isognathus, and perhaps new.
Etymology. SCHIZ = divided, ODON = teeth; ISO = even, equal,
GNATHUS = jaws.

95

96
Description
Illustrations. Fig. 25; Román 1983:98 (name misspelled as "Shizodon
sp.").
Diagnosis. This species has a distinctive pigmentation pattern
(fig. 25). It is further characterized by its bi- to pentacuspid
incisiform teeth, and a relatively short anal fin, with only seven
branched rays. Extremely similar species of Laemolvta are known from
the neighboring Capanaparo River to the south of the Apure drainage,
but they all have a superior mouth as adults, as opposed to the termi¬
nal mouth of Schizodon isognathus that have attained a size of over 40
mm SL (smaller juveniles have superior mouths and are not readily
identified).
Size. It grows to at least 300 mm SL.
Morphology. This species is slender and elongate, the greatest
body depth is slightly less than the head length. The mouth position
changes with size! Specimens examined of 20, 35, 41, 51 and 62 mm SL
have superior mouth. At 74 and 93 mm the mouth was just barely superi¬
or, and at 120 mm it was terminal.
Counts. DR ii10; AR ii7-i ii7; PR i15-i16; VR i8; LS 43-45; PDS
10-11; CPS 16; TS 10-11, 5 above, 4-5 below lateral line. Teeth 4-4 in
both jaws.
Measurements. GBD 24-27% SL.
Pigmentation. A wide black stripe extends along the midaxis of
the body from the head to the base of tail, and is accented by a pale
streak above and below. Some specimens have 3-4 dark blotches extend¬
ing vertically out from the lateral stripe. The first of these (when
present) is over the pectoral fins, the second under the dorsal fin,

97
the third between the dorsal and adipose fins, and the fourth under the
adipose fin. Juveniles less than 35 mm SL have no dark lateral band,
but rather have faint vertical bars.
Distribution and Natural Hi story
Range. It reportedly occurs in both the Amazon and Orinoco
basins, but as noted above, more than one species may be involved.
Apure distribution. Map: fig. 26. It is widespread in the
piedmont and llanos.
Habitat. It occurs in most water types, both lentic and lotic.
Abundance. COMMON.
Number of specimens examined. 296 from 92 collections.
Food. HERBIVORE. Its diet consists mainly of aquatic vegeta¬
tion, but specimens from ponds in cattle pastures were found to have
ingested manure.
Reproduction. Probable strategy: r2.
Migrations. It is known to accompany Prochilodus mariae on their
annual migrations. Large schools of fishes known locally as the ribazón,
move upstream at the onset of the dry season in November or December. Fish
remain in the highlands during the dry season, feeding on algal growth and
aquatic plants that are stimulated by the additional light they receive at
this time due to increased water clarity (lower dry-season flow reduces the
suspended sediment load dramatically). With the return of the rains,
fishes move downstream again. Many spawn at this time in midriver.
Importance. It is of only minor commercial importance because of the
numerous intermuscular bones. However, in recent years many species that
were once discarded as inedible have acquired value because of failing fish
stocks and higher prices for favorite food fishes.

Figure 26. Apure drainage distribution of Schizodon isognathus.

10°
9o
8°
f
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100
Characidae
As mentioned earl ier, because of the great diversity in their form, it
is difficult to morphologically characterize the Characidae succinctly.
Such different fishes as the slender, well-toothed pike characins, Aces-
trorhynchus; giant 40 kg herbivorous pacus or cachamas, Colossoma; a host
of small tetras such as the various species of Astyanax and Hemigrammus;
the infamous, mostly carnivorous, piranhas, Serrasalmus and Pygocentrus;
tusked, transparent scale-eating glass characins like Roeboides; salmon¬
like "dorados," Salminus, from the Andes; plate-shaped palometas, Mylos-
soma; and trout-like Brvcon all belong to this extremely heterogeneous
lineage of fishes. As taxonomic work at family and subfamily levels
progresses, some of the more widely divergent groups may be elevated to
family status or reassigned to other families.
The Characidae is the dominant family in the Apure River drainage
ichthyofauna in terms of species, absolute numbers and biomass. About 90
species were identified in the 7,642 lots (containing 149,347 individuals)
of characids examined during this study. In our study of the Apure module
fishes (Taphorn and Lilyestrom 1984), characids were always among the spe¬
cies of maximum abundance and biomass.
Knowledge of the natural history of characids is still in its infancy.
Most species are feeding generalists and can quickly switch diets to adapt
to changing food availability in the drastically different wet and dry
seasons typical of the llanos. Reproductively, all species are rl or r2,
and most are egg scatterers that provide no parental care. Larger species
tend to spawn annually with the onset of the rains. Smaller species usually
produce smaller quantities of eggs, but spawn all during the wet season, and
even into the dry if conditions remain suitable.

101
Key to the Species of Characidae of the Apure River Drainage
This key is intended only as an aid to identification of the
genera and species present in the Apure River drainage, and does not
reflect phylogenetic relationships. When appropriate, data from previ¬
ous keys have been used, including those by Eigenmann (1912, 1917),
Schultz (1944b), Géry (1972), Menézes (1976), and Machado A. (1985).
la. Dorsal-fin origin over (fig. 27a) or behind (fig. 27b) anal-fin
origin ... ...2
lb. Dorsal fin origin in front of vertical through anal-fin origin
(fig. 27c)... ...12
2a. (la) Adipose fin absent; males with elongate filament on opercle
and with lower caudal lobe elongate and sword shaped...
Corynopoma riisei (fig. 105)
2b. Adipose fin present; males lacking structures in 2a... ...3
(Note: The adipose fin may sometimes be lost through injury, so
check for stub of that fin, or examine several individuals.)
3a. (2b) Anal rays more than 55; dorsal-fin origin far behind anal-
fin origin (more than a distance equal to dorsal-fin length behind
anal-fin origin)... Xenagoniates bondi (fig. 230)
3b. Anal rays 25 to 55; dorsal-fin origin over or just slightly behind
anal origin (never more than a distance equal to dorsal-fin length
behind anal-fin origin)... ...4
4a. (3b) Maxilla longer than vertical eye diameter (fig. 28) ...5
4b. Maxilla shorter or about equal to vertical eye diameter 9
5a. (4a) Lateral line incomplete or interrupted; prepectoral region
slightly to strongly keeled... ...6
5b. Lateral line complete; prepectoral area rounded or flat 7

102
Figure 27. Relative position of dorsal and anal fin in characids.
a. dorsal-fin origin over anal-fin origin.
b. dorsal-fin origin behind anal-fin origin.
c. dorsal-fin origin in front of anal-fin origin.

103
Figure 28. Relative lengths of maxilla and vertical eye diameter.
Arrow (A) indicates length of maxilla.
Arrow (B) indicates length of vertical eye diameter.

104
6a. (5a) Fewer than 40 anal-fin rays; lateral scales 33-37; base of
anal fin scaled only anteriorly, with 1/2 to 1 scale; dorsal
midline with a black narrow stripe from occiput through dorsal-fin
base... Gnathocharax steindachneri (fig. 123)
6b. Anal-fin rays 40 or more; lateral scales 41-45; entire length of
anal-fin base sheathed with 2 or more rows of scales; dorsal
midline lacking stripe... Paragoniates alburnus (fig. 188)
7a. (5b) Upper jaw with "tusks" or out-turned conical teeth protruding
through skin of snout (figs. 29a & b); premaxilla with conical
teeth but these not elongated to form canines; maxilla not extend¬
ing back beyond middle of eye; spiny projection of cleithrum below
pectoral-fin base not very elongate, triangular or rounded in
shape and not reaching posterior edge of pectoral-fin base
(figs. 29a & b)... .. .8
7b. Upper jaw lacking "tusks" (fig. 30); premaxilla with canines;
maxilla extending back to middle of eye or beyond; spiny
projection of cleithrum (below pectoral-fin base) elongate, reach¬
ing posterior edge of pectoral-fin base (or nearly so) (fig. 30)...
Charax qibbosus (fig. 98)
8a. (7a) Upper jaw extending well beyond lower; lateral scales 67-81;
projection below pectoral notch more weakly developed, and less
strongly triangular with tip rounded; tusks stout and prominent
(fig. 29a)... Roeboides affinis (fig. 204)
8b. Upper jaw about even with lower; lateral scales 53-65; projection
below pectoral notch in cleithrum strongly triangular; tusks
moderately to weakly developed (more weakly developed in smaller
individuals) (fig. 29b)... Roeboides davi (fig. 206)

105
Figure 29. "Tusks" of Roeboides spp.
a. Roeboides affinis.
b. Roeboides dayi.

106
Figure 30. Head and anterior body of Charax qibbosus.
Note lack of "tusks" on upper jaw and long spinous projec¬
tion of cleithrum.

107
9a. (4b) Maxillary teeth present and obvious, conical in shape, and
present along entire length of bone; premaxillary and dentary
teeth conical... Heterocharax macrolepis (fig. 153)
9b. Maxillary teeth absent or not obvious, if present, conical in
shape and present only in upper corner of bone adjacent to pre¬
maxilla; premaxillary and dentary teeth tri- or multicuspid 10
10a. (9b) Body depth less than 2.5 in SL; scales ctenoid, at least in
prepelvic area; anal rays 39 to 43...
Ctenobrvcon sdí1urus (fig. 113)
(in part, see couplet 40a)
10b. Body depth more than 3 in SL; scales cycloid; anal-fin rays 28
to 35... ...11
11a. (10b) Lateral line complete; lower caudal rays in males modified
to form a spur (fig. 31); dorsal-fin origin behind anal-fin
origin; lateral scales 38-43.. .Gephyrocharax valenciae (fig. 121)
lib. Lateral line incomplete, with only 6-8 pored scales (not passing
tip of depressed pectoral fin); lower caudal rays normal; dorsal-
fin origin over anal origin; lateral scales 31-35...
Classification uncertain (fig. 232)
(Note: This fish might simply be another species of Hemigrammus,
but the dorsal fin is positioned much farther back than in most
members of that genus; furthermore, this fish has a rather flat
predorsal profile, similar to that observed in Gephyrocharax.)
12a. (lb) Snout with outward projecting "tusks" or mammila (tooth-like
projections used to scrape scales from other fishes)... ...13
12b. Snout without tusks (teeth all inside mouth)... ...14

108
Figure 31. Modified caudal rays (spur) of male Gephyrocharax valenciae.

109
13a. (12a) Color pattern unique: two very large, round black spots,
one of which covers most of caudal peduncle, the other situated on
side just anterior to dorsal fin (sometimes with a faint line
connecting these spots); lateral line complete...
Exodon paradoxus (fig. 32)
(Note: This species has not yet been collected in the Apure
drainage but occurs in the Capanaparo, Cinaruco and Meta rivers.)
13b. Color pattern not as in 13a; body without large spots, a narrow
midlateral line present, and an arrowhead-shaped, black spot
present at base of tail; lateral line incomplete...
Serrabrvcon magoi (fig. 210)
14a. (12b) Preventral abdominal midline with a series of hard, sharp-
pointed, bony serrae or scutes (sometimes only the tips emerge
from skin); body usually deep and disk-shaped... ...15
14b. Preventral abdominal midline lacking hard bony scutes and not
serrate, though the abdominal scales may sometimes form a (much
softer) keel; body shape variable... ...32
15a. (14a) Adipose-fin base length greater than or equal to distance
from posterior edge of dorsal-fin base to origin of adipose fin
(fig. 33a)... Genus Metvnnis (three spp.)... ...16
15b. Adipose-fin base length less than distance from posterior edge
of dorsal-fin base to origin of adipose fin (fig. 33b) ...18
16a. (15a) Gill rakers very long and fine, densely packed, more than
50 gill rakers on first (outer) arch; some living specimens with
red humeral spot; dorsal fin usually not spotted...
Metvnnis 1 una (fig. 169)

no
Figure 32. Pigmentation pattern in Exodon paradoxus
Figure 33. Relative adipose fin base lengths in Metvnnis spp.
a. adipose fin base length (A) greater than distance from
posterior edge of dorsal-fin base to origin of adipose
fin (B).
b. adipose fin base length (A) less than distance from
posterior edge of dorsal-fin base to origin of adipose
fin (B).

Ill
16b. Gill rakers long, but not so fine and less closely packed, fewer
than 40 gill rakers on first arch; humeral spot in living speci¬
mens usually not red; dorsal fin with or without spots... ...17
17a. (16b) Supraoccipital process relatively short, usually more than
2.6 in distance from base of occiput to dorsal-fin origin; adipose
fin not extremely long and low, the dorsal-adipose distance
usually less than 1.5 times in its base; gill rakers 17-23 on
first arch; dorsal fin usually marked with small spots...
Metvnnis argenteus (fig. 165)
17b. Supraoccipital process very long, usually less than 2.4 in dis¬
tance from base of occiput to dorsal-fin origin; adipose fin
extremely long and low, dorsal-adipose distance usually more than
1.5 in its base; about 35 gill rakers on first arch; dorsal spot¬
ted or not... Metvnnis hypsauchen (fig. 167)
18a. (15b) Dorsal rays 25 or more...Mvleus rubripinnis (fig. 183)
(Note: A juvenile, identified as M. pacu, from the Rio Aguaro,
has fin-ray counts of DR i ii20 and AR i ii31, and the incisors of
the first premaxillary row closely appressed to the second row of
molariform teeth. It is not in this key, but is described in the
species accounts, it may be confused with a juvenile Mylossoma,
and key out in couplet 31).
18b. Dorsal rays 24 or fewer... ...19
19a. (18b) Upper jaw (premaxilla) with one row of teeth... ...20
19b. Upper jaw (premaxilla) with two rows of teeth... ...29
20a. (19a) Teeth in lower jaw robust, round-based, nearly conical and
directed somewhat outward, set well apart, not touching; lower
jaw extremely evertible... Catoprion mentó (fig. 96)

112
20b. Teeth in lower jaw flattened and compressed, usually tri- or
pentacuspid, and set very close together, usually touching; lower
jaw not greatly evertible... ...21
21a. (20a) Teeth more or less symmetrically pentacuspid, the central
cusp a little larger than the other four (fig. 34a), as in most
characids, teeth not jammed together to form a single cutting edge
premaxilla and dentary with pentacuspid teeth; dorsal-fin rays 18-
20; adipose fin long, its length greater than eye diameter...
Pygopristis denticulatus (fig. 202)
21b. Teeth asymmetrically tri- to pentacuspid (fig. 34b), jammed toge¬
ther to form a single saw-toothed cutting edge; teeth of premax-
i11 a and dentary at most tricuspid, with central cusp strongest;
dorsal rays 15-17; adipose fin usually less than eye diameter...22
(Note: The section of the key which deals with the piranhas
(couplets 22-28) may not work for those individuals that have not
yet attained adult pigmentation. Juvenile pigmentation patterns
persist in most species up to about 30 mm SL. A separate key to
the juveniles is not presently possible, see Machado A. (1985) for
the identification of some juveniles.)
22a. (21b) Anal rays 29 or fewer; head and jaws wide and robust; snout
short and blunt; body stocky; dorsal profile from snout to behind
eyes convex; roof of mouth never with teeth; normal color pattern
with bright red belly and a dark black humeral spot behind opercle
(but old or reproductive individuals are often all black, and juv¬
eniles under 30 mm SL are plain with a spot on caudal peduncle as
in most piranhas); head wider, bony interorbital width less than
three times in head length...Pygocentrus caribe (fig. 200)

113
a
Figure 34. Piranha teeth.
a. Symmetrical pentacuspid teeth found in
Pyqopristis denticulatus.
b. Asymmetrically tri- or pentacuspid teeth found in most
other piranhas.

114
22b. Anal rays 30 or more; head and jaws usually more slender and
longer; snout longer; body less stocky, somewhat more compressed
than in couplet 22a; dorsal profile from snout to behind eyes
concave; roof of mouth with teeth, except in old individuals that
may wear them out; color variable, belly red only in breeding
season, if at all, pigmentation pattern (in preservative) usually
including numerous spots on sides; head narrower, bony interorbit¬
al width usually more than three times in head length... ...23
23a. (22b) Last (sixth) tooth on premaxilla low and long, its base much
wider than that of the tooth preceding it; teeth on roof of mouth
(on ectopterygoid) triangular, bi- or tricuspids, with central
cusp largest, and lateral cusps small, often hidden near base of
tooth beneath skin, these teeth flattened to form sharp cutting
edges, usually seven or more present on each side... ...24
23b. Last (sixth) tooth on premaxilla with base about as wide as that
preceding it; teeth on roof of mouth (ectopterygoid) unicuspid,
squared distally with rounded base, more conical than flattened,
usually fewer than five present on either side... ...28
24a. (23a) Caudal fin with black outer border (fig. 35c), sometimes
mostly black (fig. 35a & b)... ...25
24b. Caudal fin with a distinct, black, median or basal, vertical band;
the outer border of fin clear or hyaline (fig. 36)... ...27
25a. (24a) Caudal fin not usually black to tips, edges clear (fig.
35b); body relatively elongate, its maximum depth more than two
times in SL (without lower jaw); a silvery fish with spots on
upper sides usually running together to form vertical wavy
lines... Serrasalmus elonoatus (fig. 216)
(in part, this species also keys out in couplet 27a)

115
Figure 34. Caudal pigment patterns in piranhas.
a. caudal fin all or mostly black.
b. caudal fin mostly black with hyaline edge.
c. caudal fin with black outer edge.
Figure 36. Piranha caudal fin pigmentation showing median black band,
(basal black band not shown).

116
25b. Caudal fin all black or with black outer border (fig. 35a & c);
body deeper, its maximum depth less than two times in SL 26
26a. (25b) Body depth less than 1.7 times in SL (without lower jaw);
dorsal rays 16-19, usually 17 or more; preventral scutes 17-20;
head slender; body compressed; spots on sides frequently vertical¬
ly elongate; width of lower jaws at commissure less than diameter
of eye... Serrasalmus altuvei (fig. 212)
26b. Body depth 1.7-2.0 in SL except in largest adults (over 150 mm
SL), where it is about 1.6, (however, S. altuvei seldom reaches
150 mm); dorsal-fin rays 15-17, usually 16 or fewer; preventral
scutes 18- 24; head wide and massive; body thick; spots on sides
rounded; lower jaw very robust in largest individuals, with the
width at the commissure of the jaws is greater than the eye diame¬
ter... Serrasalmus rhombeus (fig. 220)
27a. (24a) Anal-fin rays 31 or fewer; body more elongate, its maximum
depth always more than two times in SL (without lower jaw); head
and jaws not very compressed, bony interorbital width less than
3.3 times in head length; spots on sides frequently united to form
wavy vertical lines; body and opercle usually silvery in preserva¬
tive.. Serrasalmus elongatus (fig. 216)
(in part, this species also keys out in couplet 25a)
27b. Anal-fin rays 32 or more; body less elongate, smaller specimens
sometimes have a maximum body depth that fits more than two times
in SL (without lower jaw), but body depth increases with size;
head and jaws narrow and compressed, the least bony interorbital
width more than 3.3 in head length; spots on sides rounded, seldom
united to form vertical lines; body and opercle usually tan or
yellowish in preservative... Serrasalmus irritans (fig. 218)

117
28a. (23b) Sides plain silvery with a few scattered irregular spots
(more obvious in preservative); caudal peduncle often edged poste¬
riorly with a faint, thin, black bar that extends along upper and
lower caudal-fin rays, rest of fin clear, the border not edged in
black; no humeral spot; a faint black line along base of anal fin
(fig. 37)... Pristobrvcon striolatus (fig. 198)
28b. Sides heavily spotted (more obvious in preservative); caudal
peduncle plain or with a wide, diffuse, darker central area, caudal
fin usually edged in black as in S. rhombeus (fig. 35c); a distinct
dark humeral spot just behind opercle; no black line along anal
base, although this fin often has a black outer border...
Serrasalmus medinae (fig. 214)
29a. (19b) Anal fin with fewer than 30 rays, the fin scaled only on its
base; no predorsal spine embedded in flesh immediately in front of
dorsal-fin base; body heavyset; each side of lower jaw with six or
more teeth; large species that reaches some 40 kg... ...30
29b. Anal fin with more than 35 rays, the fin scaled well beyond base;
predorsal spine present, embedded in flesh immediately in front of
dorsal-fin base (fig. 38); body disk shaped, very compressed; each
side of lower jaw with only four teeth; moderate sized species that
seldom reach over 1 kg...Genus Mvlossoma (2 spp.)... ...31
30a. Lateral-line scales usually 67-76; scale rows above lateral line to
dorsal-fin base 19-23; total gillrakers in first arch from 26 (in
juveniles) to over 146 in larger individuals; adipose-fin with
ossified rays in specimens over 55 mm SL; maxilla without teeth;
opercle very wide (fig. 39a)...Colossoma macropomum (fig. 104)

118
Figure 37. Caudal fin pigmentation in Pristobrvcon striolatus.
Figure 38. Predorsal spine found some serrasalmines.

119
30b. Lateral-line scales usually 78-90; scale rows above lateral line
to dorsal-fin base 26-30; total gillrakers in first arch 26-36;
adipose fin without ossified rays at any size; maxilla with 1-3
teeth in upper corner; opercle narrower (fig. 39b)...
Piaractus brachypomus (fig. 194)
31a. (29b) Serrae on midline of belly behind pelvic fins 18-22, the
last one very close to first anal-fin ray; branched anal-fin rays
about 37; base of adipose 2.66-3.66 in dorsal-fin base...
Mvlossoma duriventris (fig. 186)
31b. Serrae on midline of belly behind pelvic fins 10-16, the last one
not close to first anal-fin ray; branched anal-fin rays 28-34;
base of adipose fin 3.75-4.25 in dorsal-fin base...
Mvlossoma aureum (fig. 185)
32a. (14b) Anal-fin base short; anal-fin rays 16 or less... ...33
32b. Anal-fin base moderate to long; anal-fin rays 17-52... ...38
33a. (32a) Lateral line incomplete, pored scales not reaching level
of dorsal-fin origin... ...34
33b. Lateral 1ine complete... ...35
34a. (33a) Caudal fin with triangular black spot on median rays; anal-
fin rays 15-16; no thin black line on mid-ventral part of caudal
peduncle; maxilla with 5-7 tiny tricuspid teeth...
Parapristella georgiae (fig. 190)
34b. Caudal fin plain, without a spot; anal-fin rays 13-14; a thin
black line along mid-ventral part of caudal peduncle; maxilla
without teeth or with 1-2 wide teeth, each with 5-7 cusps...
Hemigrammus cf analis (fig. 129)
(in part, see also couplet 66a)

120
Figure 39. Opercular bone shape in:
a. Colossoma macropomum.
b. Piaractus brachypomus

121
35a. (33b) Scales in the row above lateral line much larger than the
pored scales of the lateral line (22 in series from opercle to
base of tail versus 32 pored lateral-line scales); dorsal-fin rays
11-12; a large species reaching over 70 mm SI
Chalceus macrolepidotus (no fig.)
(Note: This species is not yet reported from the Apure drainage).
35b. Scales along side of body all of about the same size, from 35-40
in number; dorsal-fin rays ten; smaller species seldom exceeding
70 mm SL, usually much smaller...Genus Creaqrutus (three spp.)...36
36a. (35b) Dorsal fin pigmented, ranging from a dark central black
blotch to mostly black throughout, with fin tip clear; scales in
series from lateral line to dorsal fin origin six...
Creaqrutus n. sp. (fig. Ill)
36b. Dorsal fin plain, or with a few scattered melanophores at its
extreme tip; scales in series from lateral line to dorsal-fin
origin 4-5.. .. .37
37a. (36b) Branched anal-fin rays 8-10, usually 9; predorsal scales 8-
9; adipose origin over base of last anal ray; humeral blotch
variable, but usually vertically elongate, extending well below
lateral line, and crescent shaped (opening towards head, no darker
black spot included in blotch, (fig. 40a)...
Creaqrutus bol i vari (fig. 109)
37b. Branched anal-fin rays 10-11, rarely 12; predorsal scales 10-12;
vertical through adipose-fin origin over middle of last depressed
anal-fin ray; humeral blotch rounded, sometimes extending faintly
vertically above lateral line, but rarely extending much below it,
pigment darkest in rounded spot near lateral line, the upper part
sometimes curved forward in a tight crescent, but not forming an
elongate crescent, (fig. 40b)...Creaqrutus cf beni (fig. 107)

122
Figure 40. Humeral crescents in:
a. Creagrutus bolivari.
b. Creagrutus cf beni.

123
38a. (32b) Anal-fin base long, the anal fin with 40 or more rays....39
38b. Anal-fin base shorter, the anal fin with 39 or fewer rays 43
39a. (38a) Scales cycloid, often crenate; sides often with wavy black
lines; teeth all multicuspid...Markiana geayi (fig. 161)
39b. Scales ctenoid, body rough when lightly rubbed from tail toward
head; usually no wavy black lines on sides; teeth variable 40
40a. (39b) Teeth all multicuspid, canine teeth absent; lateral-line
scales 51 or fewer... Ctenobrvcon spi1urus (fig. 113)
(in part, this species also keys out in couplet 10a)
40b. Some teeth conical, canine teeth usually present; lateral line
scales more than 51... ...41
41a. (40b) Back strongly humped, body depth usually less than three in
SL; lateral-line scales over 100; lower jaw with only one row of
teeth (no row of tiny teeth behind first row)...
Cvnopotamus bipunctatus (fig. 115)
41b. Back only slightly if at all humped, body depth usually more than
3.3 in SL; lateral-line scales 70-84; lower jaw with a second row
of tiny conical teeth behind first... ...42
42a. (41b) Black blotch usually present on chin, as well as at origin
of dorsal, anal and both lobes of caudal fins; first pair of
premaxillary canines (at symphysis) set closely to one another,
the distance between their bases is less than the basal diameter
of one tooth; branched anal rays 35-38; lateral-line scales 73-76;
ectopterygoid with ventral bony crest; nasal bone tubular...
Acestrocephalus cf boehlkei (fig. 62)
42b. No blotches of black as above, but skin along edge of lower jaw
usually with a well-defined row of melanophores along bases of

124
teeth; first canines on premaxilla not close to one another,
usually separated by a distance nearly equal to their length;
branched anal-fin rays 39-45; lateral-line scales 79-84; ectopter-
ygoid without a bony ventral crest; nasal bone laminate...
Galeocharax n. sp. (fig. 119)
43a. (38b) Lateral-line scales 54 or more... ...44
43b. Lateral-line scales 53 or fewer... ...53
44a. (43a) Teeth multicuspid, never caniniform... ...45
44b. Teeth conical, caniniform or triangular, with only weak lateral
cusps (if any)... ...49
45a. (44a) Upper jaw teeth arranged in three irregular rows; anal-fin
rays 28 or fewer; body deeper, its depth less than 3.5 in SL....46
45b. Upper jaw teeth in only two rows; anal-fin rays more than 30; body
long and slender, its greatest depth four or more in SL....48
46a. (45a) Lateral-line scales 54-56 (including those on caudal fin);
caudal fin with black crescent along its base, more extensive on
upper lobe; anal fin black along its base (fig. 41a); sides with
horizontal rows of dots but these not forming wavy lines...
Brvcon falcatus (fig. 41a)
(Note: This species is not yet reported from the Apure drainage,
but is known from Guyana and the northern Amazon Basin.)
46b. Lateral-line scales 56 or more (including those on caudal fin);
caudal fin with pigmentation as in fig. 41b or c, and always
lacking complete black crescent along its base; anal fin with or
without black along its base; posterior half of sides usually with
horizontal wavy lines... ...47

125
47a. (46b) Body with black midlateral stripe that continues out through
central caudal-fin rays to their tips in most individuals; anal
fin and base not black, the fin usually gray or clear (fig.
41b)... Brvcon whitei (fig. 84)
47b. Body without black midlateral stripe; caudal fin with asymmetrical
diagonal band of black from caudal peduncle up through basal
portion of upper lobe; anal fin with black pigment starting near
base of ninth ray and intensifying posteriorly (fig. 41c)...
Brvcon bicolor (fig. 82)
(Note: In preservative the black pigment is usually lost.)
48a. (45b) Maxilla short, barely reaching orbit in horizontal axis, and
not reaching down to ventral margin of orbit on vertical axis
(fig. 42a); teeth wide and multicuspid (with at least seven uni¬
formly-sized cusps), upper caudal lobe black on proximal half (or
more), lower lobe hvaline...Iguanodectes spi1urus (fig. 159)
48b. Maxilla long, extending back well beyond anterior edge of orbit,
and reaching down past ventral margin of orbit (fig. 42b); teeth
with central cusp much larger than others, usually tri- to penta-
cuspid, not particularly wide; both caudal lobes with wide dusky
black border... Brvconops affinis (fig. 92)
49a. (44b) Snout neither very elongate nor tapering to a point (fig.
43a); all teeth conical or triangular, uniform in size, with no
large canines standing out from rest; lateral scales 55-65; body
often with horizontal rows of dots...Salminus hi 1arii (fig. 208)
49b. Snout very elongate, narrow, and tapering to a slender point (fig.
43b); large canine teeth present which are much larger than rest
of teeth; lateral scales more than 70; body without rows of dots
forming horizontal 1ines...Genus Acestrorhvnchus (2-4 spp.) ...50

126
Figure 41. Pigmentation patterns in:
a. Brycon falcatus (not present in Apure Drainage).
b. Brycon whitei.
c. Brycon bicolor.

Figure 42. Maxilla length and shape in:
a. Iguanodectes spi1urus.
b. Brvconops affinis.
Figure 43. Snout shape in:
a. Salminus hi 1arii.
b. Acestrorhynchus spp.

128
50a. (49b) Scales very small, over 140 in lateral series and 30-38
above lateral line to dorsal-fin base; gill-rakers 30-39 on first
arch; first premaxillary tooth small, often hidden in skin near
base of first large canine situated just behind; opercle usually
with two dark patches near posterior margin...
Acestrorhvnchus falcirostris (fig. 64)
50b. Scales larger, 70-125 in lateral series and 12-22 above lateral
line to dorsal-fin base; gillrakers 15-26 on first arch; first
tooth on premaxilla a large canine; opercle usually without dark
areas... ...51
51a. (50b) Humeral region with a large black blotch...
Acestrorhvnchus falcatus (no fig.)
(Note: This species has not yet been reported from Apure drainage,
but is present in the upper Orinoco and Essequibo basins.)
51b. Humeral region without a large dark blotch, though a small one
sometimes present... ...52
52a. (51b) Anal-fin rays 25-31; lateral-line scales 108-122; scales
above lateral line to dorsal origin 20-22; maximum size larger,
reaching length of 250 mm... Acestrorhvnchus microlepis (fig. 67)
(Note: Two species may key here. The more common is the nominal
/L microlepis, the other is a very similar form with a longer
snout, called here Acestrorhvnchus sp. "pico largo," which may
prove to be a different species or perhaps the juveniles of /L
falcirostris.)
52b. Anal-fin rays 21-25; lateral-line scales 74-85; scales above
lateral line to dorsal-fin origin 12-14; a dwarf species (SL
usually less than 85 mm)... Acestrorhvnchus minimus (no fig.)
(Note: This species is not yet reported from Apure drainage, but
is present in southern Apure state).

129
53a. (43b) Canines present in both jaws, other teeth conical; opercle
with "a bite missing" from central posterior margin (fig. 44)...
Lonchoqenys ilisha (no fig.)
(Note: This species is not yet reported from the Apure drainage,
but is present in southern Apure state.)
53b. No large canine teeth present in either jaw, teeth conical to
multicuspid; opercle with margin normal... ...54
54a. (53b) Lateral line incomplete (only 4-10 pored scales present),
not reaching level of dorsal-fin origin... ...55
54b. Lateral line complete... ...72
55a. (54a) Dorsal fin with a distinct black spot or blotch; anal fin
sometimes similarly marked... ...56
55b. Dorsal and anal fins without distinct black spots, though some¬
times dusky overall or edged in black... ...58
56a. (55a) A large, well-defined humeral spot present; anal fin usually
plain, with 26-27 rays; teeth in two rows on premaxilla, the first
(outer) row consisting of one to three tiny teeth on either side,
often hidden beneath skin of upper lip...
Hyphessobrycon bentosi (fig. 155)
(Note: Some individuals, especially small ones, lack the first
row of premaxillary teeth, thus placing them in the genus Megalam-
phodus. but this character hardly seems to be of generic value.
It is possible that both genera are present, but few specimens are
available for study. Problems involved with definition of the
genera Hemigrammus, Hyphessobrycon, Megalamphodus, Moenkhausia and
Pristella are complex and beyond the scope of this work.)
56b. No humeral spot; anal fin with or without black mark and usually
with 23-25 rays; teeth in one or two rows on premaxilla 57

130
Figure 44. Anterior portion of Lonchogenys ilisha. Note the oddly
curved posterior margin of opercle which seems to have "a
bite missing".

131
57a. (56b) Premaxillary teeth all conical, long, slender and sharply
pointed, and arranged in only one row; anal fin usually with
black mark near tip of first (longest) rays...
Megalamphodus cf axelrodi (fig. 163) (See note in 57b.)
57b. Premaxillary teeth tri- to multicuspid, and arranged in two rows;
anal fin plain... Hemiqrammus cf eleqans (fig. 133)
(Note: Pristel 1 a maxillaris, a species easily confused with
Hvphessobrvcon bentosi, Mega!amphodus cf axelrodi or Hemiqrammus
cf elegans, could occur in the Apure drainage since it is found
in eastern Venezuela. However, all specimens examined so far
have lacked the toothed maxilla characteristic of P. maxillaris.)
58a. (55b) Tail with distinct, black bars on upper or both lobes
(figs. 45a, b, c)... ...59
58b. Tail without black bars on its lobes, but with a central spot
or blotch often present or tail plain... ...61
59a. (58a) Tail pattern as in fig. 45a; anal-fin rays 14-15; no
maxillary teeth... Hemiqrammus cf rhodostomus (fig. 145)
59b. Tail pattern not as above, but rather like fig. 45b or c; anal-
fin rays 20-24; 1-4 maxillary teeth present... ...60
60a. (59b) Both lobes of tail with black, as in fig. 45b...
Hemiqrammus marginatus (fig. 135)
60b. Only upper lobe of tail with black, as in fig. 45c...
Hemiqrammus sp. "arriba" (fig. 149)
61a. (58b) Caudal peduncle and tail plain, sometimes with edges or
basal area dusky, but never with a distinct midbasal spot 62
61b. Caudal peduncle and/or tail with a distinct spot or blotch
centered at base, or embedded (pigment not on surface, but deeper
in flesh) at base of each lobe... ...67

132
Figure 45. Caudal pigmentation patterns in:
a. Hemigrammus cf rhodostomus.
b. Hemigrammus marginatus.
c. Hemigrammus sp. "arriba".

133
62a. (61a) Premaxillary teeth numerous, more than seven on either side,
slender and pointed, arranged in a single row, lateral cusps weak
if present; maxilla with teeth along most of its length, more
than five in number; no black lateral midline along sides nor
other black markings; tail red or pink in life...
Aphvocharax alburnus (fig. 69)
62b. Premaxillary teeth less numerous, usually no more than five on
either side, tri- to multicuspid and arranged in two rows;
maxilla with teeth restricted to area near connection to
premaxilla, fewer than five in number, if present at all;
pigmentation on body variable, but usually with black markings
of some kind; tail red or not... ...63
63a. (62b) A tiny plain fish (usually less than 20 mm SL), only
markings are two faint embedded crescents at base of each caudal
lobe and a few scattered melanophores on tip of upper caudal lobe
(fig. 46); anal-fin rays 19-21...Hemiorammus cf mimus (fig. 141)
(Note: Hemigrammus mimus. named for its similarity to a species
of Microschemobrvcon, was described from specimens with only 17
anal-fin rays. Apure drainage specimens have 19-21.)
63b. Tiny to small fishes (adults usually 20-45 mm SL) with various
pigmentation patterns, but never with embedded crescents of
pigment base of each caudal-fin lobe; anal-fin rays 16-27 64
64a. (63b) Anal-fin rays 22-27; pored scales in lateral line 14-15,
usually reaching as far back as pelvic fins... ...65
64b. Anal-fin rays 16-22; pored scales in lateral line 8-10, not
extending much beyond the pectoral fins, if that far... ...66

Figure 46. Caudal pigmentation in Hemigrammus cf mimus.

135
65a. (64a) Anal fin with a dark black line near and parallel to its
base, strongest anteriorly, and with edge of fin often black as
well (fig. 47); humeral spot over third scale in lateral line;
anal-fin rays 22-24; caudal peduncle without red in life, although
tail may be red... Hemigrammus barrigonae (fig. 131)
(in part, see couplet 70a)
(Note: Some FL barrigonae have a complete lateral line, in which
case they will probably key to Moenkhausia copei, couplet 81a.)
65b. Anal-fin base lacking dark black line near its base, and no black
on margin of fin; sometimes with a weak first humeral spot,
followed by a second stronger spot over fifth to seventh scale in
lateral line; anal rays 25-27; caudal peduncle brilliant pink or
red in life; caudal fin either plain or red near base...
Hemigrammus stictus (fig. 151)
66a. (64b) Anal-fin rays 16 or fewer; usually only one multicuspid
maxillary tooth... Hemigrammus cf analis (fig. 129)
(in part, see also couplet 34b)
66b. Anal-fin rays 18 or more; usually two, sometimes three multi¬
cuspid maxillary teeth... Hemigrammus microstomus (fig. 139)
67a. (61b) Caudal fin with embedded crescent of black pigment at base
of each lobe (fig. 48); premaxillary teeth in one row; humeral
spot absent... Microschemobrvcon casiquiare (fig. 171)
67b. Caudal fin not pigmented as above; premaxillary teeth in two
rows; humeral spot sometimes present... ...68
68a. (67b) Caudal spot extremely large, wide and high, covering entire
caudal peduncle and much of tail (fig. 49); caudal spot weakly
continuous with black midlateral stripe...
Hemigrammus newboldi (fig. 143)
(Formerly known as Ramirezel 1 a newboldi.)

136
Figure 47.
Figure. 48. Caudal pigmentation of Microschemobrvcon casiouiare.
Figure. 49. Caudal pigmentation of Hemiorammus newboldi.

137
68b. Caudal spot smaller than above, restricted either to tail or
caudal peduncle or centered at base of tail, but never covering
most of caudal peduncle and tail; caudal spot usually not continu¬
ous with lateral stripe if latter is present... ...69
69a. (69b) Lateral line very short, with 6-7 pored scales and reaching
only to over middle of depressed pectoral fin; a wide, dusky
lateral band along sides (fig. 50), continuing through lower half
of the caudal peduncle onto central caudal rays; body heavily
pigmented in general, in many individuals the lower half of body
is black or dusky... Hyphessobrvcon metae (fig. 157)
69b. Lateral line slightly longer, with 8-10 pored scales, and usually
reaching beyond pectoral-fin tips; no wide black band as above,
although a thin midlateral stripe often present; body pigmentation
variable, but usually not with lower half more heavily pigmented
as above... ...70
70a. (69b) Body deep, its greatest depth 2.5-2.8 in SL; caudal spot
barely noticeable; black midlateral stripe present; base of anal
fin with a distinct black line, this fin often edged in black as
well (fig. 47)... Hemiqrammus barrigonae (fig. 131)
(in part, see couplet 65a)
(Note: This species is very similar to Moenkhausia copel, see
couplet 81a.)
70b. Body slender, its greatest depth 3.3 to 4.0 in SL; caudal spot
obvious, well developed (fig. 51); usually either a strong black
midlateral stripe present or a strong line present at base of
anal, but not both... .. .71

138
Figure 50. Pigmentation of Hvphessobrvcon metae.

139
a
Figure 51. Posterior pigmentation patterns in:
a. Hemiorammus cf schmardae.
b. & c. Hemiorammus micropterus.

140
71a. (70b) A distinct black line along base of anal fin (fig. 51a);
caudal spot vertically elongate, as high as caudal peduncle, with
more pigment on peduncle than on tail; pored section of lateral
line longer, reaching to over middle of pelvic fins; midlateral
stripe absent or very weak, consisting only of isolated
chromatophores; a dwarf species, the largest specimen only 25 mm
SL, and most less than 20 mm SL...
Hemiorammus cf schmardae (fig. 147)
71b. No distinct black line along base of anal; caudal spot triangular
and horizontally elongate, passing to tips of central caudal rays
in some individuals (fig. 51b), or extended vertically (fig. 51c),
in which case there is more pigment on tail than on caudal
peduncle; pored section of lateral line shorter, usually not,
sometimes barely reaching the pelvic-fin insertion; a distinct,
thin black midlateral stripe present, strongest posteriorly, but
not continuous with caudal spot; a small species, reaching about
40 mm SL... Hemigrammus micropterus (fig. 137)
72a. (54b) Lateral line normal, not strongly decurved, extending back
from opercle more or less along mid-axis of body to mid-base of
caudal fin or (in some species) slightly decurved, but never so
low as to pass within 1-4 scales of anal-fin origin, and passing
nearer to mid-axis of body (an imaginary line through eye and
midbase of caudal fin) than to anal-fin origin... ...76
72b. Lateral line strongly decurved (fig. 52), starting near upper
opercle, then curving down to within 1-4 scales of anal-fin origin,
and passing nearer to anal-fin base than to mid-axis of body....73

Figure 52. Decurved lateral line (of Triportheus sp.).

142
73a. (72b) Abdomen rounded or flat, never keeled; pectoral fins not
greatly enlarged; premaxilla with two rows of teeth; (usually only
two teeth on either side in the outer row, five in the inner row);
dentary with only one row of teeth; body anchovy-like in shape
(fig. 117)... Engraulisoma cf taeniatum (fig. 117)
73b. Abdomen in prepelvic region narrowed to form sharp-edged keel;
pectoral fins enlarged; premaxilla with three rows of teeth;
dentary with two rows of teeth (the second "row" actually just
two conical teeth behind main row near the symphysis); body not
anchovy-like in shape... genus Triportheus (three spp.) 74
74a. (73b) Scales above lateral line seven to dorsal-fin origin
(counting all scales, large or small)...
Triportheus sp. "cola roja" (fig. 228)
(Note: Color in this species is quite variable among and between
populations. The body can be plain or spotted, and the tail is
usually red, but often with a central black stripe.)
74b. Scales above lateral line to dorsal-fin origin 5-6... ...75
75a. (74b) Body relatively deep, its depth less than three in standard
length; central caudal rays usually black, often elongated;
usually six transverse scales above lateral line; body often with
rows of black dots... Triportheus anquíatus (fig. 226)
75b. Body long and slender, its depth about 3.4 in standard length;
caudal fin often yellow, with concave distal margin edged in
black usually five transverse scales above lateral line, but some¬
times a small sixth scale at dorsal origin; body plain, silvery...
Triportheus albus (fig. 224)
...77
76a. (72a) Premaxillary teeth in one row...
76b. Premaxillary teeth in two rows...
.. .78

143
77a. (76a) Premaxillary teeth usually with nine cusps, dentary teeth
usually with seven points, the central cusp larger and higher
than the rest (fig 53a); anal-fin rays 21-25 (usually 22-23); 11
transverse scale rows (from pelvic-fin insertion to dorsal-fin
origin)... Cheirodon pulcher (fig. 100)
(Note: There is still some debate as to the correct generic name
for this species, which was formerly known as Odontostilbe, Géry
(1977) used the latter name for those species with a complete
lateral line.)
77b. Premaxillary and dentary teeth usually five-pointed, the outer
cusps on each tooth minute, and the dentary teeth with three
flat central cusps of same size and height (fig. 53b); anal-fin
rays 23- 27 (usually 25-27); usually only nine transverse scales
rows... Cheirodontops geavi (fig.102)
78a. (76b) Anal-fin rays 18-25... ...79
78b. Anal-fin rays 26 or more... ...82
79a. Anal-fin rays 24-25; black line absent along anal-fin base...
Brvconamericus sp. (fig. 86)
79b. (78a) Anal-fin rays 18-23; black line either present or absent
along anal-fin base... ...80
80a. (79a) Upper caudal-fin lobe black with lighter spot near base...
Moenkhausia lepidura complex (fig. 179)
(in part, see also couplet 95b)
80b. Caudal lobes both plain, hyaline, neither black nor barred;
central rays of caudal fin sometimes dark, or with a diffuse
basal crescent...
.. .81

144
a
b
Figure 53. Multicuspid teeth of:
a. Cheirodon pulcher.
b. Cheirodontops geavi.
Teeth on the left are from the upper jaw, those on the
right are from the lower jaw.

145
81a. (80b) A distinct black line present at base of anal fin; humeral
spot horizontally elongate; anal rays 21-23; inner premaxillary
row with five teeth; maxilla with four small narrow teeth...
Moenkhausia copei (fig. 175)
(Note: This species is very similar to Hemigrammus barrigonae,
and since the latter often has a complete lateral line, they are
easily confused. The traditional character used to distinguish
Moenkhausia species, a scaled caudal fin is often unreliable and
impractical. Many Hemigrammus have partially scaled caudal fins,
and most Moenkhausia lose part or all of the caudal scales when
preserved. M^_ copei has only five rows of transverse scales
between the lateral line and the dorsal origin (vs. six or seven
in barrigonae), and has a longer snout, a more slender body,
and a more falcate anal fin.)
81b. No black line at anal-fin base; humeral spot vertically elon¬
gate; anal-fin rays 18-20; inner premaxillary row with four
teeth; maxilla with two wide, seven-pointed teeth...
Brvconamericus deuterodonoides (fig. 90)
82a. (78b) Anal-fin rays 35-37... ...83
82b. Anal-fin rays 34 or fewer... ...86
83a. (82a) Body more elongate, its greatest depth more than 2.8 in
SL; maxilla with numerous (more than eight) slender teeth;
premaxilla with small numerous (six or more) teeth in main row;
preventral area flat and with an unusual arrangement of excep¬
tionally large scales that overlap on the ventral midline and
are curved up or bent at their distal edge, small auxiliary
scales present along ventral midline (on top of larger
scales)... Phenacogaster cf megalostictus (fig. 192)

146
83b. Body deep, its greatest depth less than 2.5 in SL; maxilla with
up to four teeth, if any; premaxilla with 4-5 robust teeth on
each side in inner row; preventral area rounded to slightly flat¬
tened; abdominal scales (from between pelvic-fin insertions to
between pectoral fin origins) arranged in normal rows... ...84
84a. (83a) Caudal peduncle with very large black spot; lateral line
slightly decurved anteriorly, dropping down two or three scales
immediately behind upper opercle where it begins (fig. 54)...
Tetraqonopterus aroenteus (fig. 222)
84b. Caudal peduncle without spot; lateral line straight, not de¬
curved anteriorly... ...85
85a. (84b) Anal-fin with first rays black and often extended as
filaments, other fins plain (fig. 55); a predorsal spine embed¬
ded in flesh just anterior to dorsal origin; lateral-line scales
about 37... Poptel1 a orbicularis (fig. 196)
85b. First anal-fin rays neither black nor filamentous, all fins
often reddish in life; no predorsal spine; lateral-line scales
about 32... Gvmnocorvmbus thaveri (fig.125)
86a. (82b) Pelvic bones elongate, spine-like, and projecting anteri¬
orly through wall of abdomen in front of pelvic-fin insertions
(or visible just beneath the skin); predorsal midline partly
naked; no spot on caudal peduncle or tail; a distinct round
humeral spot present, with faint dorsal and ventral vertical
extensions, surrounded by a white area, and placed farther back
on the side than in most Astvanax, about midway between the poste¬
rior margin of opercle and dorsal-fin origin...
Astvanax polvlepis (fig. 77)
[tail scaled at base; lateral-line scales 38-42; anal-fin rays
27-29.]

147
Figure 54. Slightly decurved lateral line of Tetragonopterus argenteus.
Figure 55. Anal-fin pigmentation of Poptel1 a orbicularis.

148
86b. Pelvic bones not elongate nor projecting out through body wall
in front of pelvic-fin insertions; predorsal midline usually
normally scaled (ie. with a row of scales right down the mid¬
line), but if partly naked then caudal peduncle with distinct
horizontally elongate spot (in A_^ bimaculatusl: humeral spot
usually present, but seldom surrounded by paler whitish area,
and placed farther forward, nearer to opercle than to dorsal-fin
origin... ...87
87a. (86b) Lateral-line scales 46-54... ...88
87b. Lateral-line scales 29-45... ...89
88a. (87a) Upper caudal-fin lobe not black, caudal fin red in life,
sometimes with a black stripe through central rays (even in
faded preserved specimens a few melanophores on central caudal
rays are usually visible with magnification); a distinct double
humeral bar or spot usually present, the first vertically
elongate with a more intense ovate spot at level of pupil, the
second fainter, and also vertically elongate; maxilla short,
barely (or not) reaching posteriorly to a vertical line through
anterior margin of pupil; eye smaller, measuring more than 2.9
times in HL; BD greater, usually less than 3.0 in SL...
Astvanax integer (fig. 73)
[Predorsal series of scales normal, complete.]
88b. Upper caudal-fin lobe black with a bright orange or yellow basal
spot (whitish in preserved specimens) no humeral spot; maxilla
longer, reaching posteriorly to below center of pupil of eye;
eye large, usually measuring less than 2.9 times in HL; BD less
usually 3.5 - 4.0 in SL... Bryconops melanurus (fig. 94)

149
89a. (87b) Anal-fin base with black oblique stripe that passes through
caudal peduncle at an angle and continues up into upper lobe of
caudal fin (fig. 56). (in faded preserved specimens, asymmetrical
black pigment is usually visible at least on the tail with slight
magnification)... Astvanax metae (fig. 75)
[Lateral-1ine scales 39-41; anal-fin rays 27-33; caudal fin
scaled only near at base.]
89b. Pigmentation pattern not as in 89a... ...90
90a. (89b) Inner premaxillary row with four teeth (since this char¬
acter is somewhat variable, count the teeth on both sides of
the jaw in several specimens to be sure); second suborbital in
contact with the preoperculum below and covering cheek 91
90b. Inner premaxillary row with five teeth; second suborbital leaving
a naked area between it and lower limb of preoperculum... ...93
91a. (90a) Lateral-line scales 41-44; maxilla with up to 20 teeth
along most of its length (the number increasing with age)...
Hemibrvcon metae (fig. 127)
91b. Lateral-line scales 36-39; maxilla with 1-3 wide teeth at
proximal end... ...92
92a. (91b) Lateral stripe continuing uninterrupted through caudal
peduncle onto tail (fig. 57); anal-fin rays usually 27 to 30,
rarely 26 or 31... Brvconamericus beta (fig. 88)
92b. Lateral stripe fading out on caudal peduncle, the caudal-fin base
with faint crescent of pigment (fig. 58); anal-fin rays 24 or 25,
rarely 26 or 27... Brvconamericus sp. (fig. 86)
(in part, also keys out in couplet 79a)
93a. (90b) Lateral-line scales 33-35; both caudal-fin lobes extensive¬
ly scaled (scales often lost in poorly preserved specimens) 94

150
Figure 56.
Figure 57.
Caudal pigmentation in Astvanax metae.
Caudal pigmentation of Brvconamericus beta.
Figure 58. Caudal pigmentation of Brvconamericus sp.

151
93b. Lateral-line scales 36 or more; caudal fin at most scaled only
near base... ...96
94a. (93a) Tail plain, sometimes dusky; first humeral spot dark and
distinct, highlighted by surrounding lighter area; second humeral
bar faint and vertically elongate; body depth 2.0 - 2.1 in SL...
Moenkhausia chrvsarqyrea (fig. 173)
94b. Tail with black bars on upper, or both lobes (figs. 59 or 60);
first humeral spot faint or absent; no second humeral spot; BD
2.6 or more in SL... ...95
95a. (94b) Tail with black bars on both lobes (fig. 59a), tips of
tail often milk-white... Moenkhausia dichroura (fig. 177)
95b. Only the upper lobe of caudal fin black, tips not white (fig.
60)... Moenkhausia 1epidura complex (fig. 179)
(in part, also keys out in couplet 80a)
(Note: This is very similar to M^ dichroura in morphometric and
meristic characteristics, and perhaps may only represent local
variation in color pattern and not distinct species.)
96a. (93b) Predorsal midline partially to completely naked, the
scales from either side sometimes irregularly overlapping or
touching, but the row down the midline mostly absent...
Astvanax bimaculatus (fig. 71)
[Humeral spot distinct, horizontally ovate. Caudal peduncle
with horizontally elongated spot that extends weakly onto
mid-caudal rays.]
96b. Predorsal midline with normal series of scales... ...97

152
Figure 59.
Caudal pigmentation
in Moenkhausia dichroura.
Figure 60. Caudal pigmentation in Moenkhausia lepidura complex.

153
97a. (96b) Sides of body with distinct, black, wavy ( < shaped) lines
(fig. 61); humeral spot horizontally ovate, distinct, not
usually vertically elongate; up to six teeth on proximal end of
maxilla... Astvanax superbus (fig. 61)
97b. No wavy lines on sides of body; humeral spot vertically elongate
and often diffuse; only 1-2 teeth near proximal end of
maxilla... Astvanax venezuelae (fig. 80)
Figure 61. Astvanax superbus.

154
Species Accounts
Acestrocephalus cf boehlkei Menézes 1977
Dientón Alargado - Pike Characin
Fig. 62. Map: fig. 63. Couplet 42a.
Generic synonymy
Acestrocephalus Eigenmann 1909b:316 (type species: Xiphorhamphus anomalus
Steindachner 1879a:48; Eigenmann 1910:447; Menézes 1976:37 (synonymy).
Specific synonymy
Acestrocephalus boehlkei Menézes 1977:185-193, fig.l (type locality: Rio
Punino, trib. to Río Payamino [Napo River drainage, Amazon River
Basin], above Coca, Ecuador), (seen).
Types. Holotype: MCZ 51473. Paratypes: MCZ 51474 (2), 51475 (1);
MZUSP 12993-94 (2), 12995 (1),12996-98 (3); USNM 216140 (1), 216141 (1);
ANSP 134500 (5).
Comments. Only three species of Acestrocephalus are known: A. anoma-
1 us (Steindachner 1879), A. sardina (Fowler 1913b) and A. boehlkei Menézes
1977. The nominal A. boehlkei is known from isolated upper tributaries of
the Amazon in Brazil and the Rio Ventuari drainage in the upper Orinoco
Basin in Venezuela. Comparison with the literature shows that Amazonian
specimens differ slightly in pigmentation and some of the counts from speci¬
mens from the Apure drainage. More specimens and a direct comparison with
the types are necessary to determine the true status of the Apure drainage
population, which probably represents a new species.
Etymology. ACESTRO = a thick needle, referring to the canine teeth,
CEPHALUS = head; BOEHLKEI = named to honor the late Dr. James Bohlke, a
North American ichthyologist who dedicated many years of study to South
American fishes.

155
Figure 62. Acestrocephalus cf boehlkei.

156
Description
Illustrations. Fig. 62; Menézes 1977: fig. 1, photo of paratype.
Diagnosis. This species is distinguished from Galeocharax sp., the
most similar species in the Apure drainage, by the presence of black
pigment on the chin and at the origins of the dorsal and anal fins when
small. Adult specimens can be distinguished by having fewer branched
anal-fin rays (35-38 vs 39-45) and fewer lateral-line scales (73-76 vs.
79-84). It is further distinguished in having the first pair of
premaxillary canines (at the symphysis) very closely set, instead of
separated by a distance nearly equal to their length in Galeocharax sp.
See Menézes (1977) for additional osteológica! characters that separate
Acestrocephalus from Galeocharax. This species differs from the other
cynopotamine characids of the Apure drainage in lacking the
characteristically "humped" dorsal profile.
Size. It reaches about 150 mm SL, but most individuals are between
80-100 mm.
Morphology. This fish is a slender (BD more than 3.3 in SL) medium¬
sized cynopotamine characid, with ctenoid scales and a smoothly sloping
dorsal profile that is not much more curved than the ventral. The snout is
elongate but not greatly narrowed, the maxilla is prominent and heavily
toothed (30-45). The dentary has a second row of tiny conical teeth behind
the first. The cleithrum is not notched.
Counts. DR ii9; PR i15; VR i7; LLS 73-76.
Pigmentation. Most small specimens have a black dot or blotch at the
tip of the chin, at the base of the dorsal and anal fin origins, and at the
base of the upper and lower caudal lobes. These are lost in most adults.
The body is light brown in alcohol, but silvery in life, with a faint

157
humeral spot and a weak, blackish lateral stripe that is strongest
behind the level of the dorsal fin, and widens to form a faint blotch
on the caudal peduncle. The fins are all hyaline, but the caudal fin
has a black crescent at its base.
Pistribution and Natural History
Range. This species is known from the Río Ñapo Rio Payamino and
Río Coca en Ecuador, the Rio Tulumayo in Peru (Menézes 1977).
Apure distribution. Map: fig. 63. A. boehlkei is known only
from the lower portion of the Apure drainage. Although it is taken
infrequently, the localities are well separated and encompass the
entire lower drainage.
Habitat. It occurs near shore in the main river channel of the
Apure and other large lowland rivers, often in or near aquatic vegeta¬
tion.
Abundance. UNCOMMON. Only 13 collections at just a few localities
produced specimens of this species, but since it inhabits main river chan¬
nels where sampling was seldom undertaken during this study, its abundance
is probably greatly underestimated by the above figures.
Number of specimens examined. 80 from 13 collections.
Food. CARNIVORE. It is probably mainly piscivorous.
Reproduction. Probable strategy: rl. Though this species spawning
habits are unknown, small (20-30 mm SL) juveniles are present from May and
September, indicating that it spawns throughout the wet season, and perhaps
all year long.
Migrations. Since this species inhabits the main river channels,
long migrations are not expected, but it probably makes local movements
to avoid low water conditions during the dry season.

Figure 63. Apure drainage distribution of Acestrocephalus cf boehlkei.


160
Acestrorhvnchus falcirostris (Cuvier) 1819
Care’perro, Picúa - Pike Characin
Fig. 64. Map: fig. 65. Couplet 50a.
Generic synonymy
Acestrorhvnchus Eigenmann & Kennedy 1903:527 (new name for Xiphorhynchus
Agassiz, preoccupied in Aves, Swainson 1827, and for Xiphorhamphus
Muller & Troschel preoccupied in Aves, Blyth 1843); Eigenmann 1903:146
(type species: Salmo falcatus Bloch 1794:120, by subsequent
designation of Eigenmann 1910:447); Menézes 1969:33 (diagnosis,
synonymy).
Xi phorhvnchus Agassiz 1829:76 (type species: Sal mo fal catus Bloch,
by subsequent designation of Jordan & Evermann 1917:132).
Xiphorhamphus Miiller & Troschel 1844:92 (type species: Salmo fal catus
Bloch, by subsequent designation of Jordan 1919:221).
Sphvraenocharax Fowler 1906:460. (type species: Xiphorhamphus abbreviatus
Cope 1878:687, by original designation).
Specific synonymy
Hvdrocyon falcirostris Cuvier 1819:361 (type locality: Brazil), (seen).
Xi phorhvnchus falcirostri s Agassiz 1829:76; Cuvier & Valenciennes
1849:341.
Hvdrocyon armatus Schomburgk 1841 pi. 25 (not the description).
Xiphorhamphus falcirostris Müller & Troschel 1844:92 (Brazil:
description).
Acestrorhvnchus falcirostris Fowler 1906:462; 1914:254 (British Guiana);
Fernández Yépez 1955:3 (Venez.); Menézes 1969a:219 (synonymy, diagno¬
sis, Venezuelan specimens cited from Guariquito and Chimire rivers,
Guárico state) Mago L. 1970:69; Géry 1977:320 (diagnosis); Menézes &
Géry 1983:590 (key); Román 1985:162 (diagnosis).

161
Figure 64. Acestrorhvnchus falcirostris. Drawing from Menézes (1969).

162
Comments. More specimens must be examined to be certain that this
species is the same as that reported for the Amazon drainage. Menézes
(1969b) reported no significant differences in body proportions among
specimens he examined from the Amazon, Orinoco and Essequibo rivers, but
did note a trend to an increase in number of scales from north to south
and a few minor count differences. He had very little comparative
material.
It is possible that the following "species" listed as A. sp. "pico
largo" (meaning long snout in Spanish) are merely juveniles of this
species, but they differ in counts and dentition, and no similar adults
were found at the site where they were taken.
Etymology. ACESTRO = a thick needle, referring to the canines,
RHYNCHUS = jaws; FALCI = sickle shaped, ROSTRIS = snout.
Description
II1ustrations. Fig. 64; Menézes 1969b:figs. 43, 61f; Géry
1977:320. Román 1985:162.
Diagnosis. This species is distinguished from other Acestrorhvnchus
by the presence of a small conical tooth, sometimes hidden under a flap of
skin, located anteriorly to the first large canine of the premaxilla. It is
also somewhat more elongate and slender than most species. The maximum body
depth measures 5.9-6.2 times in the SL (Géry 1977). In general appearance
it is otherwise similar to A. microlepis.
Size. Although reported to reach 400 mm SL (Géry 1977), specimens of
that size are probably scarce in the wild.
Counts. DR Ü9; AR vl8-24; PR 14-19; VR 8; 140-175 scales in the
lateral line; 30-37 scales from lateral line to origin of dorsal fin, 17-22
from lateral line to origin of anal fin; GR 30-39.

Figure 65. Apure drainage distribution of Acestrorhynchus falcirostris.


165
Pigmentation. The body is silvery in life, tan in alcohol, and
darker dorsally. Fin membranes are hyaline or clear but the rays of the
dorsal fin (especially the first three), caudal fin and anal fin are
gray to black. The base of the caudal fin has a small, nearly round
black blotch that extends onto the caudal peduncle. The opercle is
edged with black posteriorly and may also have two dark concentrations
of pigment, the upper separated from the lower by a narrow light stripe
(Menézes 1969b).
Pistribution and Natural History
Range. This species is known from the Brazilian Amazon, the
Orinoco Basin in Venezuela, and Guyana (Menézes 1969b).
Apure distribution. Map: fig. 65. A. falcirostris is known only
from one site in a blackwater river of the Aguaro River system in the
southeastern corner of the drainage.
Habitat. This species was found in a nearly transparent tea-colored
stream of the low llanos of Guárico state, an area characterized by
sandy soils and sparse savanna vegetation.
Abundance. RARE. Known only from one specimen.
Number of specimens examined. 1.
Food. CARNIVORE. This species is mostly piscivorous.
Reproduction. Strategy: Probably r2.
Acestrorhvnchus sp. ("pico largo")
Care’perro, Picúa - Pike Characin
No Figure. Map: fig. 66. Couplet 52a.
Comments. The number of lateral-line scales apparently increases
with overall size in A. mi crolepis. If the same is true for A.
falcirostris (which has a LLS count 140-175) then these specimens may

166
simply represent juveniles of that species. However, the only specimen
of A^_ falcirostris collected was taken from similar blackwater habitat
in Guárico state. Since the collecting effort at the site where these
small specimens were found was extensive (monthly for one year plus
several subsequent visits), and no adults of A. falcirostris were ever
captured, their exact identification remains uncertain.
Description
Diagnosis. This species is very similar to juvenile A. microlepis
but has a longer snout.
Size. Only small specimens up to about 50 mm SL were collected.
Counts. Lateral-line scales: 99-124.
Measurements. The snout is longer than in A^ microlepis for
specimens of comparable size, hence the "pico largo" designation, which
means "long snout" in Spanish.
Pigmentation. The body is tan in preservative, with a black spot
near the base of the caudal fin, similar to that in A. microlepis or A.
falcirostris.
Pistribution and Natural Hi story
Apure distribution. Map: fig. 66. This species is known only from
one site, Caicara Creek, in northern Apure state.
Habitat. It occurs in blackwater creeks of low llanos.
Abundance. RARE. It is known only from two collections at the
same site.
Number of specimens examined. 37 from 2 collections.
Food. CARNIVORE. Presumably this species is almost exclusively
piscivorous, as is true of related species.

Figure 66. Apure drainage distribution of Acestrorhynchus sp. "pico largo


169
Reproduction. Probably r2. The juveniles collected were taken in
June and July, suggesting that breeding occurs during the early wet
season, or perhaps just prior to it.
Acestrorhvnchus microlepis (Schomburgk) 1841
Care’perro - Pike Characin
Fig. 67. Map: fig. 68. Couplet 52a.
Specific synonymy
Xi phorhamphus mi crolepis Schomburgk, 1841:247 (not plate 25; type
locality: Essequibo River), seen.
Xiphorhvnchus microlepis Cuvier & Valenciennes 1849:342 (Guyana).
Acestrorhvnchus microlepis Eigenmann 1910:447; 1912:408 (part from
Guyana); Eigenmann & Allen 1942:276 (synonymy); Fernández Y. 1955:4;
Menézes 1969a:219 1969:62 (diagnosis, redescription, distribution,
synonymy); Mago L. 1970:69; Géry 1977:330 (key); Menézes & Géry
1983:590 (key); Nico & Taphorn 1986:794 (diet).
Acestrorhvnchus cachorro Fowler 1939:274 (type locality: Rio Ucayali
drainage, Boca Chica, Peru).
Xiphorhvnchus falcatus (not of Bloch 1794:120); Cuvier & Valenciennes
1849:337 (part, Mana?).
Acestrorhvnchus falcatus Roman 1985:163 (Venez.).
Comments. This species apparently grades into A. quianensis, a form
said to be present in easternmost Venezuela and Guyana, and it is probable
the two are distinct only at the subspecific level. Alternatively, the two
could be nearly identical sister-species, sympatric throughout their range
(see above).
The holotype of A. quianensis (FMNH 74353) has jaws of approximately
equal length. The first canine tooth of the upper jaw is larger than

Figure 67. Acestrorhynchus microlepis. From Menézes (1969).

171
second. The caudal spot is triangular, and although no humeral spot is
evident, there are a few chromatophores behind the opercle. The lateral¬
line canals have lateral branches, contrary to what was indicated in Géry’s
(1977) key.
Etymology. MICRO = small, LEPIS = scales.
Description
Illustrations. Fig. 67; Menézes 1969b:67, fig.50; Géry 1977:317.
Román 1985:163, (misidentified as A. falcatus).
Diagnosis. This species is distinguished from A. falcatus by the
absence of a large humeral blotch. It has fewer lateral-line scales (108-
122) than the very similar A. falcirostris but more than A. minimus (74-85).
Menézes (1969b) gave a complete diagnosis.
Size. A. microlepis grows to about 250 mm SL.
Morphology. This is a long slender, nearly cylindrical fish, with
elongate well toothed jaws.
Counts. DR Ü9; AR v25-31; PR 13-18; PR 8; 108-122 pored scales in the
lateral line; 20-22 scales from 1 ateral line to origin of dorsal fin, 15-18
from lateral line to origin of anal. The lateral-line scale count seems to
increase with the size of the specimen, small ones around 100-115, larger
ones with up to 140. This character is quite variable. Since it is quite
unusual for the number of lateral-line scales to vary with size, it is
possible that there are two sympatric species present, a smaller one with
fewer scales and a larger one with more scales. If this is the case, they
are nearly identical. Gillrakers 20-26 on lower half of the outer arch.
Pigmentation. This fish is silvery in life, (tan in alcohol), with
the dorsum darker than the ventrum. There is a small, rounded, distinct
black spot at the base of the central caudal-fin rays. The humeral spot is

172
usually absent, or there may be a very small indistinct dark blotch at
the origin of the lateral line, behind the opercle.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins, as well as in
Guyana, Surinam and French Guiana (Menézes 1969b).
Apure distribution. Map: fig. 68. This species was most fre¬
quently collected from northern Apure state, the Aguaro River system in
the southeasternmost part of the drainage, and in Barinas in the Suripá
drainage.
Habitat. It is found most commonly in northern Apure state, in
blackwater creeks, and prestamos of the module system, and in the
tributaries of the Aguaro River system in the sandy-soiled savannas
where morichales are common. This species seems to prefer slow-moving
streams with abundant aquatic vegetation.
Abundance. COMMON, but it is found only in isolated populations where
conditions are favorable.
Number of specimens examined. 334 in 46 collections.
Food. CARNIVORE. This species is piscivorous in all but the
smallest juvenile stages, when aquatic invertebrates are taken (Nico &
Taphorn 1985).
Reproduction. Probable strategy rl. Small juveniles have been
collected in October, which is the end of the rainy season. This
probably is an indication that this species spawns more than once
during the wet season.
Importance. Although A. microlepis has no commercial importance
as a food fish, it is sometimes collected as an ornamental. A notori¬
ous piscivore, this species should be housed by itself in aquaria.

Figure 68. Apure drainage distribution of Acestrorhynchus microlepis.

-p>

175
Aphyocharax alburnus (Gunther) 1869
Sardinita Coliroja - Bloodfin Tetra
Fig. 69. Map: fig. 70. Couplet 62a
Generic synonymy
Aphyocharax Giinther 1868b:245, 254 (type species: A. pusi 11 us Gunther
1868); Eigenmann 1910:429; 1912:312; 1919:22; Eigenmann & Allen
1942:265.
Specific synonymy
Chirodon al burnus Giinther 1868:424 fig. 2 (type locality: Rio Marañon
[Peruvian Amazon Basin]), not seen.
Aphiocharax (error) alburnus Eigenmann & Eigenmann 1891:55.
Aphyocarax (error) alburnus Ulrey 1895:292.
?Aphiocarax (error) (Chirodon) alburnus Perugia 1897:25 (Rio Beni).
Aphyocharax alburnus Eigenmann 1910:429; 1915b:27 (Brazil, Bolivia),
1919:27; Román 1985:166 (Venez.).
Aphyocharax avary Fowler 1913b:532 fig.8 (type locality: Rio Madeira).
Comments. The distinctions between this species and A. ervthrurus are
slight, and they may be synonyms. Both have been cited for the Orinoco
Basin (Fowler 1943 [Florencia, Colombia] & Schultz 1944b [Caripito,
Venez.]). The Apure specimens have melanophores along the central caudal
rays, a feature stated to be diagnostic by Eigenmann (1919) since the cen¬
tral caudal rays of A. ervthrurus are said to be pale. They al so have 15-18
teeth on the maxilla, so in these two characters they agree better with the
descriptions of A. alburnus. Since A. al burnus is the older name, it would
have priority if they are synonyms. However, I have not observed the
"golden crown" coloration mentioned as characteristic of A. alburnus by
Axelrod et al. (1971), so doubt remains. The holotype (FMNH 53579) of

176
Figure 69. Aphvocharax alburnus.

177
A. ervthrurus has no humeral spot nor any dark pigment remaining, and
in general appears to be more slender than the Apure specimens, with an
apparently longer head and caudal peduncle and a larger eye; the LL is
incomplete with about 11 pored scales, the base of the caudal fin is
scaled, and the maxilla is very large and wide.
Etymology. APHYO = small, CHARAX = pal isade of pointed sticks, from
the Greek (Eigenmann 1919), in this case referring to a palisade of pointed
teeth, but also given to mean a "gnasher of teeth" in some of Eigenmann’s
other works, Charax is a characid genus frequently used in the formation of
generic names; ALBURNUS = whitish, from the Latin "albus," perhaps refer¬
ring to the whitish fin tips, or the generally light colored body.
Description
Illustrations. Fig. 69; Fowler 1948:171 fig. 196; Román 1985:166; as
A. ervthrurus: Axelrod et al. 1971:F-48.10; Géry 1977:349.
Diagnosis. There is only one species of Aphvocharax known from the
Apure drainage. Characteristic features of this species include: a single
row of conical teeth on the premaxilla; a toothed maxilla; an incomplete
lateral line and a brick-red caudal fin.
Size. A. alburnus can grow to about 50 mm SL, but most individuals
are usually between 20-40 mm. A 28 mm specimen SL weighed 4 grams.
Morphology. This is an elongate, cylindrical species (GBD is about
26% SL) with the dorsal fin well in advance of the anal fin, the pelvic fins
are in advance of dorsal; an adipose fin is present; the caudal fin has
somewhat rounded lobes. The snout is rounded and short, its length is less
than the eye diameter. The lateral line is incomplete, with 10-11 pored
scales. The base of the caudal fin is densely scaled. Males have hooks on
the pelvic and anal fins.

178
Counts. DR ii8-ii9; AR i i15-iii18; PR i9; VR i7, GR 10; LS 35-39
(with 10-11 pored scales in the lateral line reaching to just over the
pelvic insertions); TS 10; CPS 14. The premaxilla has 7-8 teeth on either
side in a single row. The maxilla has up to about 20 teeth, but is not
completely toothed, and does not reach the large suborbital posteriorly.
Pigmentation. The body is bluish-silvery to olive-tan, with the
caudal fin bright to brick red, and the anal fin with the first few rays
whitish. All fins may have some reddish tint. The body is tan in preserv¬
ative with all the fins clear (the red is usually lost). A diffuse humeral
spot is usually present. The peritoneum is gray.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map; fig. 70. This species is ubiquitous in the
llanos and Andean piedmont.
Habitat. This species survives in a wide variety of habitats in
the lowlands and Andean foothills, from major river channels to stag¬
nant ponds, and in all types of water, white, black and clear. It
usually is found in large schools, near the edges of stands of aquatic
vegetation.
Abundance. ABUNDANT.
Number of specimens examined. 6,992 from 323 collections.
Food. OMNIVORE. It feeds on aquatic and terrestrial insects,
microcrustaceans and other small invertebrates, as well as some vegeta¬
ble matter.
Reproduction. Strategy: rl; fecundity: 617 eggs/female; egg diameter:
0.65 mm. Breeding mainly occurs during the first three months of the wet
season, but perhaps continues throughout its duration. Individuals can

Figure 70. Apure drainage distribution of Aphyocharax alburnus.

00
o

181
breed more than once a year. Generation time is about 11-12 months (Wine-
miller & Taphorn 1989). Hoedeman (1975) reported on the breeding behavior
of a similar species, A. rubripinnis, in which mating takes place when
mature fish move into the shallows, and eggs and sperm are released over
aquatic vegetation in mass spawning bouts. The parents may eat the embryos.
After hatching, the embryos absorb the yolk sac for 3-4 days before begin¬
ning active foraging for plankton.
Importance. Ornamental.
Astvanax bimaculatus (Linnaeus) 1758
Sardina Dos Puntos - Two-spotted Astyanax
Fig. 71. Map: fig. 72. Couplet 96a.
Generic synonymy
Astyanax Baird & Girard 1854:26 (type species: Astvanax argenteus, by
monotypy); Fowler 1948:35.
Poeci1urichthvs Gill 1858:414,417 (type species: Poeci1urichthvs
brevoortii Girard, by subsequent designation of Jordan 1919:281).
Zygogaster Eigenmann 1913:23 (type species: Zyqoqaster fi 1iferus
Eigenmann, by original designation).
Specific synonymy
Sal mo bimaculatus Linnaeus 1758: 311 (type locality: "in America
meridionali"), seen.
Tetragonopterus maculatus Müller & Troschel 1845:14, pi.3, fig. 4 (type
locality: Brazil, Guiana, Surinam); Peters 1877:472 (Calabozo,
Venez.); Pellegrin 1899:157 (Apure River, Venez.); Regan 1906:384
(Venez.).
Charax bimaculatus Gray 1854:154 (Surinam).
Astyanax bimaculatus Eigenmann 1920:11 (near Maracay and Valencia,

182
Figure 71. Astvanax bimaculatus

183
Venez.); Pearse 1920:19 (Valencia area, Venez.); Fowler 1931:408
(Guanoco area, Venez.)? 1948:38 (synonymy, subspecies, Brazil);
Schultz 1944b:355 (Rio Guárico and tributaries, Caripito, Venez., key,
synonymy); Mago L. 1970:69.
Comments. Detailed analysis of regional variations will probably
reveal a complex of species and subspecies that are currently all
lumped.
Etymology. ASTYANAX was a Greek mythological figure, the son of
Hector. This genus was so named because of the large silvery scales’
resemblance to armor (Innes 1966); BI = two; MACULATUS = spotted.
Description
111ustrations. fig. 71; Géry 1977:412, 413, 419; Hoedeman 1975:638;
Román 1985:171; Axelrod et al. 1971:F-92.00.
Diagnosis. This species is distinguished from the other
Astyanax present in the Apure drainage in having: the predorsal midline
partially to completely scaleless; a dark black, horizontally ovate
humeral spot; a dark black, horizontally ovate caudal spot at the base
of the central caudal-fin rays that can extend onto the caudal peduncle;
and 35-42 lateral-line scales. The sides lack dark wavy lines, and
there is no oblique black stripe from the base of the anal fin onto the
tail.
Size. Although literature reports claim it can reach nearly 160 mm
TL, Apure drainage specimens reach a maximum of about 90 mm SL, and most
individuals are only 50-60 mm.
Morphology. This is a robust, deep-bodied Astyanax. The lateral
line is complete. The mid-dorsal line in front of dorsal fin is at
least partly naked, but there are often a few scales present near

184
dorsal fin. The dorsal-fin length is about equal to head length; the
anal margin is straight in adults, but emarginate in young. The swim
bladder is two-chambered, the posterior chamber nearly twice as long as
the anterior.
Counts. DR ii9; AR 29-35; PR i 11-i13; LLS 35-42; scales above LL:
7-9; scales below LL: 6-7; 9 upper and 14 lower rakers on first gill
arch; premaxillary teeth 4-4 in the outer row, and 5-5 in the inner;
principal dentary teeth 4-4; no maxillary teeth.
Pigmentation. The body is silvery to bronze, with the back
sometimes olive-green to greenish-bronze. There is a dark rounded
humeral spot behind the upper part of the opercle, over the third to
sixth or the second to fifth scale of the lateral line (Eigenmann 1912),
and sometimes accompanied by a ventrally projecting vertical streak
(Schultz 1944b). There is another dark black streak on caudal peduncle.
A blackish lateral stripe usually unites these two spots. In life the
smaller, more slender male sometimes has red or yellowish areas in the
dorsal and caudal fins. The peritoneum is black.
Pistribution and Natural History
Range. Various subspecies have been described from Panama in the
north to the La Plata drainage in southern South America.
Apure distribution. Map: fig. 72. This species is well
distributed throughout the drainage with the exception of the highest
mountain streams.
Habitat. Astvanax bimaculatus is found in almost all biotopes,
but is more common in flowing water.
Abundance. ABUNDANT.
Number of specimens examined. 6,704 from 319 collections.

Figure 72. Apure drainage
distribution of Astyanax bimaculatus.

CUENCA DEL RIO APURE
• 4MC4
00
en

187
Food. OMNIVORE. Aquatic plants, small seeds, flowers and various
aquatic and terrestrial insects, spiders, and filamentous algae comprise the
diet of this species (K. Winemiller pers. com., and original data). Pearse
(1920) reported that in Lake Valencia A. bimaculatus feeds on a wide variety
of foods including: algae (27), plants (25), insects (18), fish (11), chi-
ronomid pupae (3.9), cladocera (1.8), Diptera (1.5), Sialis larvae (0.5),
insect larvae (0.3), Gasteropoda (0.2), unidentified (5.9); the numbers
reflect per cent volume for 17 fishes examined.
Reproduction. Strategy: r2; generation time: about 12 months; fecun¬
dity: 4287 eggs/female; egg diameter: 0.90 mm (Winemiller & Taphorn 1989).
This species breeds annually during the first 2 months of the wet season.
In captivity it is easily spawned, but parents must be provided with a good
diet. They scatter eggs over aquatic plants and then abandon them.
Hatching occurs within 24-36 hours: after 5 days the yolk-sac is absorbed
and the fry become free swimming and ready to feed on plankton (Axelrod et
al. 1971).
Vanzolini & Reboucas (1965), who studied growth rings in Brazilian
populations of this species, found that up to four different annul i could be
formed in a year, and that their specimens reached adult size (about 110 mm)
in one year.
Importance. Ornamental.
Astvanax integer Myers 1930
Sardina Coliroja - Red Tailed Astyanax
Fig. 74. Map: fig. 74. Couplet 88a.
Specific synonymy
Astvanax integer Myers 1930:67 (type locality: Guaicaramo, Rio Guavio in
the Rio Meta drainage, Colombia); Fowler 1943:238 (Villavicencio,
Colombia), seen.

188
Figure 74. Astvanax integer.

189
Astvanax abramoides Schultz 1944b:355 (key, from Río Guárico and tribs.;
synonymy).
Tetragonopterus abramis Pellegrin 1899:157 (Apure River, Venez.).
Types. Holotype: CAS-SU 23726.
Comments. Following Schultz (1944b), I had originally identified
this species as A. abramoides a species described by Eigenmann from
British Guiana, but I now believe it is closer to A. integer Myers.
The holotype of A. abramoides (FMNH 52863) is a very deep-bodied fish,
the depth in SL about 2.5, and it has an ovate humeral spot just behind
the opercle on a level with the upper 1/2 of the eye. The caudal fin
is marked with a basal blotch, and the middle caudal-fin rays show
traces of pigment. The eye is quite large. There is no trace of a
lateral stripe. The dorsal midline is not scaled. There are about
seven short gill rakers on the upper limb of the outer gill arch, and
about ten longer ones on the lower arch. The teeth are quite robust,
with five tricuspid teeth on the outer premaxillary row, five pentacus-
pids on the inner row. Their rounded edges face in opposite direc¬
tions.
Specimens from the Apure drainage differ greatly from the type
material of A. abramoides. They have the dorsal midline scaled regu¬
larly, lack the caudal blotch, and in general fit the description of A.
integer much better.
Etvmoloqy. INTEGER = complete, whole or unbroken, is from the
Latin, and although no explanation was specifically indicated by the
describer, I suspect that this refers to the complete scalation of the
dorsal midline observed in this species, a character that is of
subgeneric importance in these fishes.

190
Description
II1ustrations. Fig. 74; Fowler 1943:236, fig. 17; Román
1985:173, as A. abramoides.
Diagnosis. When alive, this species is easily recognized by the
bright red tail. It usually has more lateral-line scales (43-50,
usually 46-50) than other Apure drainage Astyanax, although some over¬
lap exists. A. polvlepis sometimes has high lateral-line scale counts,
but that species is characterized by stout pelvic spines that often
project through the abdominal skin, a situation never observed in A.
integer. It is easy to confuse this species with A. bimaculatus. but
that species lacks maxillary teeth (one is present in A. integer, and
has 43 or fewer lateral-line scales (vs 43-50).
Size. It is a fairly large Astyanax reaching at least 120 mm SL.
Morpholoqv. See Myers (1930) for a complete description. His
diagnosis is as follows: Astyanax with predorsal region completely and
regularly scaled, lateral line about 50, anal 30, the central caudal
rays black, a well defined humeral blotch, and a wide dark lateral band
from behind humeral spot to caudal. Fowler (1943) described in detail
two specimens from Villavicencio, Colombia.
Counts. DR ii8-ii9; AR 1Ü26-Í1131; PR 11-13; VR i7; LLS 43-50,
usually 46-50; TS 18 (9-10 above LL, 7-8 below it). Premaxillary teeth
3-4 or 4-4 in outer row, 5-5 in inner. Maxillary with one tooth.
Scales at base of caudal not covering more than 1/4 of the length
longest rays. Gillrakers about seven on upper and 10-14 on lower limb
of outer arch.
Measurements. GBD about 38% SL; HL about 28% SL.
Pigmentation. The body is silvery with a vertically elongate,
often double humeral spot. In life the caudal fin is bright red, with

191
the central caudal rays sometimes blackish. The anterior anal rays have
red along their shafts and black specks on the posterior membranes.
There is sometimes red or yellow color in the dorsal fin and eye. A
wide lateral stripe starts under the tip of the last dorsal ray and
widens noticeably on the caudal peduncle; it is silvery in life, but
dark in preservative. It is usually not continued to the tips of
central caudal-fin rays. The adipose fin is speckled with pigment.
The humeral spot extends ventrally, and is not usually horizontally
ovate.
Distribution and Natural History
Range. It occurs in Andean tributaries of the Orinoco River.
Apure Distribution. Map: fig. 74. This species occurs in Andean
montane and piedmont rivers and tributaries of the Guariquito River in
the easternmost region of the drainage.
Habitat. This species is mostly restricted to piedmont and montane
streams, though it has also been collected from the main channels of
rivers in the low llanos, and from the upper reaches of the blackwater
streams in open savanna. It is usually found in clear, clean,
relatively undisturbed streams with moderate to fast currents, over
rock, gravel or sand substrates. It is abundant in pools and the calmer
parts of the rapids, and often takes refuge behind branches or boulders
where it if often seen schooling.
Abundance. COMMON.
Number of specimens examined. 691 from 106 collections.
Food. OMNIVORE. Its diet includes small seeds, fruit and terres¬
trial insects (Winemiller, pers. com.). This species is a midwater
feeder that picks food items from the debris carried downstream in the

Figure 74. Apure drainage distribution of Astyanax integer.


194
current. It will eat all sorts of campsite leftovers (rice, spaghetti,
fruit, bread, meat etc.), an indication that it is not too particular
about its diet, scavenging whatever the current washes down.
Reproduction. Strategy: r2; generation time: 12 months; fecundity:
8400 eggs/female; egg diameter: 1.00 mm; oviparous; it spawns annually
during the first two months of the wet season (Winemiller & Taphorn
1989).
Importance. Ornamental.
Astvanax metae Eigenmann 1914
Sardina - Astyanax
Figs. 56 & 75. Map: fig. 76. Couplet 89a.
Specific synonymy
Astvanax metae Eigenmann rn Eigenmann, Henn & Wilson 1914:11 (type
locality Rio Negro, Villavicencio, Colombia), seen; Eigenmann 1920:11
(Maracay and Valencia area, Venez.); Pearse 1920:19,42 (Lake Valencia
area, Venez.); Eigenmann 1922:235 (Lake Valencia area, Venez.) Schultz
1944b:358 (Río Guárico and tribs, and Caripito, Venez.); Mago L.
1970:69; Géry 1977:397; Román 1985:173.
Types. Holotype: FMNH 56640 (formerly CM 5457), (examined).
Paratypes: CAS 39229 (4) (formerly IU 13153).
Etymology. METAE = for the Río Meta.
Description
111ustrations. Figs. 56 & 75; Géry 1977:397; Román 1985:173.
Diagnosis. The oblique band of black pigment that runs along the
base of the anal fin into the upper lobe of the caudal fin is
characteristic of this species.
Size. This is a large Astyanax reaching over 160 mm TL, but
usually seen between 80-100 mm TL.

195
Figure 75. Astvanax metae.

196
Morphology. A. metae is a large, robust Astvanax with a moderately
compressed body that is rather heavy anteriorly and which has the
preventral area rounded. The predorsal midline is completely scaled,
with a series of 10-11 scales. The lateral line is complete. The
dorsal-fin origin is about an eye diameter closer to the snout tip than
to the midcaudal base, or equidistant between them (Schultz 1944b). The
occipital process extends about one-fourth of the distance to the dorsal
fin origin, and is bordered by four scales on each side. The maxilla is
slightly longer than the eye diameter. The anal and caudal-fin bases
are sheathed with scales. The teeth are massive, and number four in the
outer and five in the inner row on each side of the premaxilla. There
are four large teeth plus 3-6 tiny teeth on either side of the dentary.
Counts. DR ii9; AR 27-33; PR i11-i14; LLS 39-42; TS above LL 9-10,
below LL 6-7; GR 14-17; premaxilla with 4-5 teeth in outer and 5 in
inner row; maxilla with 1 or 2 teeth; GR 12 on upper and 15 on lower
1imb of outer arch.
Measurements. GBD 2-2.6 in large adults, 2.9 in young; HL about
29% SL in adults.
Pigmentation. The body is silvery, the fins clear. An oblique
black band extends posteriorly from the base of the anal fin up onto
the caudal peduncle and thence onto the upper middle caudal-fin rays.
The black pigment is distributed asymmetrically. It is most intense on
the four central rays of the upper caudal-fin lobe, with practically no
black pigment on the (opposite) central rays of the lower caudal-fin
lobe. This oblique stripe is sometimes lost in largest specimens, or
those long preserved. The humeral spot is absent or represented by an
oblique streak, with a pale area behind it (Eigenmann 1914, Schultz

Figure 76. Apure drainage distribution of Ast.yanax metae.

UD
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199
1944b). The holotype still shows the characteristic black streak along
the anal base and into the tail, with the caudal rays above center the
darkest, and black to their tips.
Pistribution and Natural Hi story
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 76. This species is limited to the
highlands.
Habitat. Mountain streams with clean water swift currents, and
rocky substrates are the favorite haunts of this species. It often
forms mixed schools with A. integer. Hemibrvcon metae and other
characids the congregate below riffles and behind submerged branches.
Abundance. COMMON.
Number of specimens examined. 1,002 from 70 collections.
Food. OMNIVORE. These fish feed on items washed down in the swift
current such as small seeds, fruit, terrestrial insects (Winemiller
pers. com.). Pearse (1920) reported a diet in Lake Valencia consisting
of 45% (by volume) mayfly nymphs, plants (46%), and seeds (6%).
Reproduction. Strategy: r2; generation time: 12 months; fecundity:
9528 eggs/female; egg diameter: 1.00 mm; it spawns annually during the
first month of the rainy season (Winemiller & Taphorn 1989).
Astvanax polvlepis (Giinther) 1864
Sardina - Astyanax
Fig. 77. Map: fig. 78. Couplet 86a.
Specific synonymy
Tetragonopterus maculatus Miiller & Troschel 1845:74 (part).
Tetragonopterus polylepis Giinther 1864:320 (type locality: Guyana), seen.
Poecilurichthys polylepis Eigenmann 1910:433; 1912:356.

200
Figure 77. Astvanax polvlepis.

201
Etymology. POLY = many; LEPIS = scales.
Description
Illustrations. Fig. 77; Eigenmann 1912:pi. 52.
Diagnosis. The circular spot set farther back on the sides (over
the seventh to the ninth scales of the LL) distinguishes this species
from other Apure drainage Astyanax species. The black spot is
surrounded by a pale area, and may have vertical extensions. Spines
that extend out from the pelvic bones (and at least in preserved
material, pierce the skin of the abdomen) are another feature unique to
this species among Apure drainage Astvanax.
Size. Normally A. polvlepis is found between 40-50 mm SL, but it
reportedly reaches 90 mm (Eigenmann 1912).
Morphology. The body is much compressed and elevated, more than
in any other Apure drainage Astyanax. There are 3-4 scales on the
dorsal midline just in front of the dorsal-fin origin, but anteriorly it
is naked. A sheath of scales is present anteriorly on the anal fin
and the base of the caudal fin is also scaled.
Counts. DR ii9; AR iii23-iii26; PR i11-i12; VR i7; LLS 38-46; TS
9-10 above, 7-8 below LL. The premaxillary teeth number four in the
outer row, five in the inner, and the maxilla has two multicuspid teeth.
Measurements. GBD 36-41% SL; HL 27-32 % SL.
Pigmentation. The body is whitish in preservative, but heavily
dotted with melanophores, especially along the lateral line area and the
dorsal midline. The scales on the upper sides are outlined with gray
and have white centers. Pigment in the dorsal fin follows right next to
the rays. The anal fin has melanophores throughout the membrane and
sometimes the first rays are tipped in white.

Figure 78. Apure drainage distribution of Astyanax polylepis.

CUENCA DEL RIO APURE
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204
Distribution and Natural History
Range. It occurs in Guyana and Venezuela.
Apure distribution. Map: 78. A. polvlepis is most commonly taken
from blackwater streams in northern Apure state, from the Aguaro River
system in the southeasternmost part of the drainage, and in Barinas
state, in the Suripá drainage.
Habitat. Blackwater streams and morichales are the most often
frequented biotopes of this species.
Abundance. COMMON.
Number of specimens examined. 642 from 41 collections.
Food. OMNIVORE.
Reproduction. Strategy: Probably r2, as are the rest of the species
of Astvanax. It probably scatters eggs over vegetation, and no parental
care is expected. Males have tiny hooks on the anal fin rays presumably
used to assist in maintaining close contact during the fertilization of
the eggs.
Importance. Ornamental.
Astvanax superbus Myers 1942
Sardina - Superb Astyanax
Fig. 61. Map: fig. 79. Couplet 97a.
Specific synonymy
Astvanax superbus Myers 1942:92 fig.2 (type locality: a small brook
tributary to Rio Tamanaco [a trib. to Río Paye, Río Portuguesa
system], at Camoruco, 20 km northeast of San Carlos, [state of
Cojedes], Venezuela); Schultz 1944b:357 (key, Rio Torbes near Táriba,
Venez.); Mago L. 1970:69; Géry 1977:430,454 (key).

205
Comments. Dr. Barry Chernoff (pers. com.) believes this species
might pertain to the genus Astvacinus.
Types. Holotype: CAS-SU 36489, Paratypes: CAS-SU 36490 (3).
Etymology. SUPERBUS = we cannot know what Myers considered to be
"superb" in this fish, since the significance was not explained, but I
suspect that it refers to the intricate color pattern of wavy lines, an
unusual situation for this genus.
Description
Illustrations. Fig. 61 (in key) ; Myers 1942:92 fig. 2 (drawing
of holotype).
Diagnosis. The dark wavy lines on the sides of this species are
unique among Apure drainage Astvanax species. It also has a rather
high number of maxillary teeth (six). The humeral spot is dark and
horizontally ovate. A dark lateral stripe continues out through the
central caudal-fin rays.
Size. It grows to 130 mm SL, but most individual are smaller
(70-100 mm).
Morpholoqv. The predorsal midline is irregularly scaled, but
there is no naked area. A. superbus is a fairly typical Astvanax with
a rounded belly, the postventral area sharper, and the predorsal area
bluntly keeled. However, the body is not particularly compressed. The
dorsal-fin origin is midway between the snout tip and the tip of ap-
pressed adipose fin. The origin of the anal fin is slightly posterior
to the base of last dorsal ray, its border is nearly straight. The
appressed pelvic fins reach slightly past the anus in males, but not
quite to the anus in females. The pelvic fin insertions are just
anterior to the dorsal fin origin. The pectoral fins reach the pelvic

206
fins in males but not in females. The anal fin has a sheath of one row
of scales.
Counts. DR 10; AR 29-32; LLS 39-42; TS 7-8 above LL, 6-7 below it;
GR-12 on lower limb of first arch; maxilla with 5-6 teeth.
Measurements. Head 4.1 - 4.6 in SL, depth 2.7; eye 3.3 in a small
male to 4.1 in a large female.
Pigmentation. The body is dusky in general, darker dorsally,
with all scales except those on the belly with a narrow dark border.
The sides of the body have horizontal wavy lines between the scale
rows. Males are darker and show a more complex pattern of wavy lines
than females. A rounded or horizontally ovate humeral spot is present,
but is not surrounded by a lighter area as in A. polylepis. The later¬
al stripe is not particularly dark except on the caudal peduncle, but
the pigment continues through the central caudal rays out to their
tips.
Pistribution and Natural History
Range. It is known only from Venezuela, but is undoubtedly present
in the Andean tributaries of the Orinoco of Colombia as well.
Apure distribution. Map: fig. 79. Restricted to montane and piedmont
streams.
Habitat. Small, usually very shallow, shady streams in the Andes.
Abundance. UNCOMMON. This species may be more common that the
number of collections would indicate, since it lives in the higher areas
of the drainage, which are less accessible.
Number of specimens examined. 278 from 23 collections.
Food. OMNIVORE. It feeds on terrestrial insects and small seeds
(K. Winemiller pers. com.)

Figure 79. Apure drainage distribution of Astyanax superbus.

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00

209
800 eggs/female; egg diameter: 0.65 mm; spawning occurs annually during
the first month of the wet season. Individuals may spawn more than once
(Winemiller & Taphorn 1989).
Importance. Though not currently sold as such, this species has
potential as an ornamental.
Astvanax venezuelae Schultz 1944
Sardina - Venezuelan Astyanax
Fig. 80. Map: fig. 81. Couplet 97b.
Specific synonymy
Astyanax venezuelae Schultz 1944b:359 fig. 54 (type Locality: Rio Torbes,
1 km above Táriba, Orinoco System, [Venezuela], seen); Mago L.
1970:69; Géry 1977:442.
Types. Holotype: USNM 121449; Paratypes: USNM 121450.
Etymology. VENEZUELAE = after the country where it was collected.
Description
Illustrations. Fig. 80; Schultz 1944b:360 fig.54.
Diagnosis. This species is similar to A. superbus but lacks the
wavy lines on the sides characteristic of that species and has fewer
(1-2) maxillary teeth. The predorsal midline is scaled. It differs
from A. fasciatus in having the dorsal-fin origin much farther forward.
Size. It grows to a maximum of about 120 mm SL, but most indi¬
viduals are smaller (70-100 mm SL).
Morphology. It is a stout, somewhat compressed Astyanax.
Counts. DR ii9; AR 26-29; PR i13-i14; LLS 40-41; TS 8-9 above, 6-
7 below LL; GR1 5-16; outer premaxillary row with 4-5 teeth, inner with
five; maxilla with one tooth; all teeth tri- to pentacuspid with the
central cusp longest.

210
Figure 80
Astvanax venezuelae.

Figure 81. Apure drainage distribution of Ast.yanax venezuelae.

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213
Measurements. HL 26-27 % SL; GBD 36-37% SL.
Pigmentation. The humeral bar is distinct and vertically elon¬
gate, with pale areas in front and behind. The lateral stripe is
faint, but intensifies on the caudal peduncle and continues to the tips
of the central caudal rays. The body is silvery on the lower sides,
but the upper sides and back are brownish. The outer tips of the
anal-fin rays are black. The inside of the operculum as well as the
peritoneum are blackish.
Pistribution and Natural Hi story
Range. It is probably present throughout the Andes in the Orinoco
Basin in Venezuela and Colombia.
Apure distribution. Map: fig. 81. To date it has been taken only
in the western portion of the Apure drainage, in piedmont and montane
streams and rivers.
Habitat. Fast-flowing montane streams with clear water and rocky
substrates are the principal habitat of this species.
Abundance. UNCOMMON.
Number of specimens examined. 105 from 10 collections.
Food. OMNIVORE. This species undoubtedly feeds in a similar
manner to A. metae and A. Integer, taking seeds, fruits, insects, and
other items that are washed down with the current.
Reproduction. Strategy probably r2. Spawning is probably annual
and synchronized with the onset of the rainy season.
Brvcon bicolor Pellegrin 1909
Palambra, Bocón - Brycon, South American Trout
Figs. 41c & 82. Map: fig. 83. Couplet 47b.

214
Figure 82. Brycon bicolor.

215
Generic synonymy
Brycon Müller & Troschel 1844:90 (type species: Brycon falcatus Müller &
Troschel 1844:90, by original designation.)
Chalcinopsis Kner 1863:226 (type species: Brycon striatulus Kner &
Steindachner in Kner 1863:226, by original designation.)
Megalobrycon Günther 1869:423 (type species: M. cephalus Günther 1869,
by original designation).
Catabasis Eigenmann & Norris 1900:349 (type species: C. acuminatus
Eigenmann & Norris 1900, by original designation).
Bryconodon Eigenmann 1903:33 (type species: Brycon orthotaenia Günther
1864, by original designation).
Othonophanes Eigenmann 1903:145 (type species: Brycon 1abiatus
Steindachner 1879:75, by monotypy.
Triurobrycon Eigenmann 1909:33 (type species: Brycon 1undii Reinhardt
1874, by monotypy).
Hoiobrycon Eigenmann 1909:33 (type species: Brycon pesu Müller &
Troschel 1845:16,30, by original designation).
Specific synonymy
Brycon bicolor Pellegrin 1909:12 (type locality: Orinoco), not seen;
Myers & Weitzman 1960:103; Mago L. 1970:69; Géry 1977:339 (key); Howes
1982:11 (review of genus, synonymy).
Brycon falcatus Román 1985:164 (Venez.).
Types. Paratypes MNHN 87-746-748 (111,114.5 and 118. 5 mm SL).
Comments. The identification of the Apure drainage specimens is
still in doubt, though the form matches the description given by Howes
(1982) fairly well, the color pattern seems to be variable, and quite
similar for several species described from or reported from Venezuela.

216
Other possibilities could be Brvcon lonoiceps Steindachner 1879,
which was reported for Venezuela from Ciudad Bolivar by its describer
as well as by Eigenmann and Eigenmann (1891), who probably were merely
repeating the earlier citation. B. coauenani Steindachner 1915 was
described from the Coquenan River in Venezuela.
Etymology. BRYCON = "gnasher of teeth" from the Greek, according
to Howes (1982), but Eigenmann (1903) gave a different meaning, "to
devour" and yet another ("greedy eater") in 1927; BICOLOR = of two
colors, probably referring to the silver body with black markings.
Description
111ustrations. Figs. 41c & 82; Howes 1982:13 fig.8 (drawing of
holotype). Román 1985:165, (photo misidentified as B. falcatus).
Diagnosis. The color pattern readily distinguishes this species
from EL whitei (see figs. 41c and 82).
Size. This species grows to at least 250 mm SL.
Morphology. The premaxillary teeth are in three rows, as in all
Brvcon. The first row has 8-10 teeth, the second 8-9, and the third
only two. The lateral line is complete. Howes (1982, Table 5) gave
measurements of the three paratypes.
Counts. DR 11; AR 26-28; PR i13; VR i7; LLS 55-62; TS 15 above, 9
below LL; maxillary teeth 23-27; inner dentary teeth 18-22; GR 14-16.
Pigmentation. A dark band extends obliquely from the base of the
caudal peduncle to across the upper lobe of the caudal fin (much as in
Astvanax metae). The pigment on the anal fin begins at about the base
of the ninth ray and widens to cover the remainder of the fin posteri¬
orly. Howes (1982) considered this color pattern fins to be diagnostic
for this species, however, the black pigment is quickly lost in pre¬
servative.

217
Distribution and Natural Hi story
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 83. This species is known only from
the eastern portion of the drainage, in Guárico state, from the sandy
soils and black waters of the Aguaro River system.
Habitat. It frequents Clearwater rivers with sandy or rocky
substrates, as well as blackwater streams and large rivers, and
morichales that drain the sandy-soiled savannas of the easternmost
portion of the Apure drainage.
Abundance. RARE.
Number of specimens examined. 3 from 3 collections.
Food. OMNIVORE. B. bicolor probably feeds on all sorts of small
animals, fishes, fruits and seeds, as do most members of the genus. I
have observed this species while diving. The fishes stayed in an ill-
defined school, roving from one end of the pool to the other.
Reproduction. Strategy: r2. Spawning is probably annual and
synchronized with the onset of the seasonal rains.
Migrations. Though this species is not definitely known to do so,
it probably migrates in conjunction with the wet-dry season
changeovers, as do many similar species of Brvcon present in Brazil
and Argentina.
Importance. This species is a food fish of native peoples, and an
increasingly popular sport fish with spear fishermen and hook and line
anglers. It is seldom seen in the commercial catch because they are not
easily obtained in large numbers.

Figure 83. Apure drainage distribution of Brycon bicolor.

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Brycon whitei Myers & Weitzman 1960
Palambra - South American Trout
220
Fig. 41b & 84. Map: fig. 85. Couplet 47a.
Specific synonymy
Brycon whitei Myers & Weitzman 1960:99, fig. 1 (type locality: vicinity
of Los Micos, north end of the Cordillera Macarena, -3° 20’ N 73° 56’
W, headwaters of the Rio Guaviare (just south of Villavicencio,
Colombia), seen; Mago L. 1970:69; Cala 1977:1; Géry 1977: 335,339;
Howes 1982:46; Lilyestrom & Taphorn 1983:53 (biology).
Types. Holotype: CAS-SU 48818; Paratypes: CAS-SU 44817.
Etymology. WHITEI = after General Thomas D. White who collected
the type specimens.
Description
Illustrations. Figs. 41b & 84; Myers & Weitzman 1960:100. Géry
1977:325, (as unidentified Brycon from Rio Aguaro, Venezuela); Román
1985:165; Lilyestrom & Taphorn 1983:55, 57.
Diagnosis. The presence of a continuous black lateral stripe
from the opercle to the tips of the central caudal-fin rays readily
distinguishes this species from B. bicolor.
Size. Specimens of at least 450 mm SL were once fairly common, but
these days usually younger, smaller specimens are seen (about 250 mm
SL). Pollution and deforestation have taken a heavy toll.
Morphology. See Myers and Weitzman (1960) for a complete
description. The upper jaw teeth are arranged in three or four rows,
(depending on how you define the rows). This species is slender when
young, but as it matures individuals become increasingly heavier and
deeper bodied.

221
Figure 84. Brycon whitei.

222
Counts. DR 11; AR iii21-iii22; LLS 66-70 (given as 66 + 4 or 5 in
description).
Measurements. GBD 3.1-3.2 in SL; HL 4.0-4.2. The length - weight
curve for this species is log P = -1.73 + 3.049 Log L (n = 68, r =
0.933) P is the weight in grams and L is the standard length in cm.
Pigmentation. The body is silvery to golden, darkest dorsally,
and lighter on the sides with the belly whitish. The wide, black lateral
stripe is prominent, and continues out onto the central caudal rays. A
faint humeral spot is located just above the origin of the lateral
stripe. The pectoral and adipose fins are gray in preservative, but in
life these fins (especially the caudal) have yellow to red tints.
Pistribution and Natural History
Range. Orinoco Basin in Colombia and Venezuela.
Apure distribution. Map: fig. 85. The palambra is found in
montane, piedmont and upper llanos streams in the Andes, and in the
upper reaches of the Aguaro River system.
Habitat. This species inhabits mountain, piedmont and high
llaneran streams and rivers. It prefers clear waters, with little or no
sediments, strong currents, and high levels of dissolved oxygen. The
water temperature is usually between 24 and 28° C, with the pH nearly
neutral. It is seen in both pools and rapids, but is more common where
the gallery forest that provides most of the species’ diet is intact.
The substrate is sand, gravel or rock in their favored haunts. It is
usually seen in groups, rarely alone.
Abundance. UNCOMMON. Since this species easily avoids nets, its
true abundance may be greater than indicated by our collections. It is
still fairly common in Andean streams not yet affected by deforestation

Figure 85. Apure drainage distribution of Brycon whi

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CUENCA DEL RIO APURE
UNIVERSIDAD NACIONAL EXPERIMENTAL
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"EZEOUIEL ZAMORA"
VICE-RECTOR ADO EN PORTUGUESA
MIHIGA A

225
or pollution. In the upper Rio Caparo in 1980, this species was still
quite abundant; however, that river has since been dammed.
Number of specimens examined. 42 from 24 collections.
Food. OMNIVORE. Seeds and terrestrial insects comprised the diet
of specimens examined by K. Winemiller (pers. com.). It readily accepts
Astvanax integer used as bait, indicating that small fishes are also
taken. Lilyestrom & Taphorn (1983) found 82% of 68 specimens dissected
to contain vegetable material (flowers, leaves and seeds from trees),
with only 11.3% containing animal remains, including caterpillars,
grasshoppers, fish and a lizard.
Reproduction. Strategy: r2. Generation time: 24 months, fecundity:
over 170,000 eggs/female; egg diameter 1.5 mm. Spawning is annual, and
occurs during the first month of the wet season (Winemiller & Taphorn
1989). Lilyestrom & Taphorn (1983) found specimens in February (middle
dry season) with gonads just beginning to mature.
Migrations. It is known to migrate extensively up (at the
beginning of the dry season, Oct.-Dec.) and downstream (with the onset
of the rains in May to June).
Importance. An excellent and highly sought sport fish, also
important in subsistence fishing, this species is currently in danger of
local extinction. It has already been eliminated from large areas due
to habitat loss, alteration and pollution.
Brvconamericus sp.
Sardinita - Tetra
Figs. 58 & 86. Map: fig. 87. Couplets 79b & 92b.
Generic synonymy
Brvconamericus Eigenmann i_n Eigenmann, MacAtee & Ward 1907:139 (type
species: B. exodon Eigenmann 1907, by monotypy).

226
Figure 86. Bryconamericus sp.

227
Knodus Eigenmann 1910:216 (type species: Knodus meridae Eigenmann 1911,
by monotypy).
Comments. The pigmentation pattern of this species doesn’t
agree well with that of any described Bryconamericus. It probably
represents a new species.
Etvmology. BRYCON = is from the Greek for "gnasher of teeth"
according to Howes (1982), but Eigenmann (1903) gives a different
meaning, "to devour" and yet another "greedy eater" in 1927; AMERICUS =
from the Americas, South America in this case.
Description
Illustrations. Figs. 58 (caudal pigmentation) & 86.
Diagnosis. This species is distinguished by its low number of
anal-fin rays (usually 24-25, but up to 27). B. beta usually has 27-30
rays, rarely 26 or 31, and B. deuterodonoides has a maximum of 18. It
is further distinguished from B. beta by not having the lateral stripe
continued through the central caudal-fin rays, as is characteristic of
that species.
Size. This fish seldom exceeds 50 mm SL, and most individuals are
usually between 30-40 mm.
Counts. DR 10; AR 24 (18), 25 (22), 26 (8), 27 (2).
Pigmentation. The body is plain silvery in life, sometimes with a
faint lateral stripe. The fins are colorless. Pigment at the base of
the caudal fin forms a faint vertical crescent.
Distribution and Natural History
Range. Orinoco Basin.
Apure distribution. Map: fig. 87. This species is most common in
the low llanos, but also ranges into the mountains.

Figure 87. Apure drainage distribution of Bryconamericus sp.

229

230
Habitat. It inhabits Whitewater rivers, and is more common in the
main river channels than the other two species.
Abundance. COMMON.
Number of specimens examined. 3,096 from 77 collections.
Food. OMNIVORE. This species diet consists of seeds, insects, and
microcrustaceans.
Reproduction. Strategy: probably rl. It probably spawns in the
shallows of the rivers and the flooded plains during the annual floods.
Bryconamericus beta Eigenmann 1914
Sardinita - Tetra
Fig. 57 & 88. Map: fig. 89. Couplet 92a.
Specific synonymy
Bryconamericus beta Eigenmann in Eigenmann, Henn & Wilson 1914:7 (type
locality: Villavicencio, Colombia), seen; Eigenmann 1920:11 (Concejo,
Rio Tiquirito, Venez.); 1922:236 (northern Venezuela); 1927:389, pi.
91, fig.2 (key, redescription); Schultz 1944b:340; Mago L. 1970:69.
?Bryconamericus sp. Eigenmann 1920:11 (Lake Valencia, Venez.).
Knodus beta Géry 1977:395.
Types. Holotype: FMNH 56648 (formerly CM 5465a), examined.
Paratypes: FMNH 56649 (formerly CM 5466a-d), CAS 39492(4) (formerly IU
13158) all examined.
Comments. The form present in the Apure drainage is nothing like
the types of B. beta, but traditionally has been identified as such. It
has several more lateral-line scales than the types of B. beta and is
similar to B. caucanus (in Géry [ 1977] it will key out to that
species), but the latter is deeper bodied and supposedly limited to the

231
Figure 88. Brvconamericus beta.

232
Cauca drainage in Colombia. It is also similar to B. osgoodi from the
upper Amazon drainages of Peru, but that species is known only from the
type. There seems to be little to differentiate B. beta and B. alpha,
both described by Eigenmann from Villavicencio, Colombia. A careful
revision of material from all of the Andean headwaters of the Orinoco
will be necessary to unravel this problem.
Etymology. BETA = second.
Description
Illustrations. Figs. 57 (caudal stripe) & 88; Eigenmann 1927:389,
pi. 91, fig. 2.
Diagnosis. The lateral stripe continues onto the central caudal
rays in this species, as opposed to the situation in Brvconamericus s£.
Though there is some overlap, B. beta beta usually has 27-31 anal rays,
vs 24-26 in B. sp. B. deuterodonoides has only 17-18 anal-fin rays.
Size. This species reaches about 50 mm SL, but most individuals
are between 30-40 mm.
Morphology. The anal fin has a sheath of scales anteriorly and the
caudal fin is scaled on only about 1/4 of the lower lobe. The lateral
line is complete. The branched rays of the pelvic fins in males have
hooks.
Counts. DR ii8; AR 26(2), 27(9), 28(14) 29(13) 30(10) 31(4); PR
12; VR i7; LLS 34-38, TS 12; premaxilla with 4-4 teeth, maxilla with
two.
Pigmentation. The humeral spot is vertically elongate but vague in
many specimens. The lateral stripe is continuous with the humeral spot,
but faint anteriorly. It widens on the caudal peduncle and extends out
onto the central caudal rays, but not usually to their tips. All fins

Figure 89. Apure drainage distribution of Bryconamericus beta.

CUENCA DEL RIO APURE
OCAL* IMflCt
ro
co
4^

235
have some peppering of pigment. The dorsal scales are outlined with
pigment and there is a concentration of pigment between the pelvic fins.
Distribution and Natural History
Range. Orinoco Basin.
Apure distribution. Map: fig. 89. This species is most common in
the upper part of the Apure drainage, though scattered lowland
localities have been found.
Habitat. This species occurs in upland rivers with moderate to
fast currents and rocky substrates, as well as in smaller shaded streams
in the mountains.
Abundance. COMMON.
Number of specimens examined. 2,838 from 140 collections.
Food. OMNIVORE. Their diet includes terrestrial insects, small
seeds, and aquatic insects (K. Winemiller, pers. com.)
Reproduction. Strategy: rl; fecundity: 796 eggs/female; egg
diameter: 0.95 mm; generation time: 4 months. This species probably
spawns throughout the year. Individuals may reproduce more than once a
year (Winemiller & Taphorn 1989).
Bryconamericus deuterodonoides Eigenmann 1914
Sardinita - Tetra
Fig. 90. Map: fig. 91. Couplet 81b.
Specific synonymy
Bryconamericus deuterodonoides Eigenmann i_n Eigenmann, Wilson & Henn
1914:7 (type locality: Villavicencio, Colombia, Rio Negro [quebrada]),
seen; 1927:389 pi.91 fig.2; Fowler 1943:240; Géry 1977:387 (key).
Bryconamericus deuterodonoides deuterodonoides Schultz 1944b:340 (key);
Inger 1956:435 (key).

236
Figure 90. Bryconamericus deuterodonoides

237
Types. Holotype: B. deuterodonoides FMNH 56644 (formerly CM 5461),
examined. Paratypes CAS 61221(6) (formerly III 13156), examined.
Comments. Schultz (1944b) described B. d. eurvodous from the Rio
Guárico and tributaries between San Sebastián and San Casimiro, and
from Caripito, Venezuela (types USNM 121468).
The holotype of B. d. caudovittatus Inger (FMNH 45706) has a black
streak through central caudal rays, four teeth in inner premaxillary
row, 2-3 in outer, and two very wide maxillary teeth.
The holotype of B. d. deuterodonoides (FMNH 56644), from Vi 11avi-
cencio, Colombia in the Meta river drainage has three large, wide teeth
on the maxilla, with four teeth in the inner and 2-3 in the outer
premaxillary row. It has the base of the tail scaled. It has neither
a lateral stripe, nor any other pigment remaining. It is very similar
to Venezuelan specimens, but has a differently shaped head.
The subspecies described by Schultz (1944b) and Inger (1956) hardly
seem justified unless further distinctions can be demonstrated.
Schultz (1944b) himself questioned the difference between B.
cismontanus Eigenmann 1914 and B. deuterodonoides. both described in
the same paper along with B. alpha and B. beta, all four from the same
locality at Vi 11avicencio, Colombia. Thus, four sympatric species of
Brvconamericus are described from the Andean piedmont of the Meta
drainage. Since I have found three species in the Apure drainage, this
is perhaps possible, but it will take comparison of fresh collections
from Vi 11 avicencio with the Apure material to make final determina¬
tions.
Etymology. DEUTERODON = Deuterodon is a genus of characid, OIDES =
like; referring in this case to the similar teeth.

238
Description
Illustrations. Fig. 90; Eigenmann 1927: 389 pi. 91 fig. 3 (photo
of types).
Diaqnosis. B. deuterodonoides has only 17-18 anal-fin rays.
This immediately distinguishes it from other Brvconamericus in the
Apure drainage, since those species have 24 or more. The subspecies B.
d. eurvodous Schultz 1944b, and B. d. caudovittatus Inger 1956 are
reported to have four teeth in the outer row of the premaxilla, versus
only two in B. d. deuterodonoides, and they allegedly differ in
branched anal-fin ray counts and gill-raker number (Inger 1956).
Size. It grows to about 50 mm SL, but most individuals are
smaller, 30-40 mm.
Morphology. This species is more oval in cross section, and not as
compressed as B. beta. It has a modified, elongate axillary scale, and
a complete lateral line.
Counts. DR i i8; AR Ü15-16; PR 12; VR i7; LLS 34-35; TS 10;
premaxillary teeth 4-4, maxilla with three, dentary with nine. Inger
(1956) reported six upper and 9-10 lower gill-rakers on the outer arch
in the holotype and paratypes of B. d. deuterodonoides.
Pigmentation. The humeral spot is dark, with a pale area in
front and behind it. There is no caudal spot. The lateral stripe
begins just behind the humeral spot (which it does not touch) and
continues onto the tail, but fades before reaching the tips of the
central caudal rays. Flecks of black pigment in the dorsal, adipose
and anal fins are concentrated distally. The dorsum has numerous black
specks, but the dorsal midline is darkest. The nose, lips and chin are
flecked with melanophores, and the cheeks are white with red dorsally.

239
The eye is black below the pupil. The upper base of the caudal fin has
a characteristic red to yellow dot in life.
Distribution and Natural History
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 91. It is mostly limited to Andean
piedmont and montane streams.
Habitat. This species is found in upland, white and Clearwater
streams and rivers, usually over rocky substrates.
Abundance. COMMON.
Number of specimens examined. 5,187 from 117 collections.
Food. OMNIVORE. Its diet includes aquatic insects, small seeds,
and terrestrial insects.
Reproduction. Strategy: rl; fecundity: 243 eggs/female; egg
diameter: 0.85 mm; generation time: about 6 months. Individuals
probably spawn more than once a year (Winemiller & Taphorn 1989).
An extremely unusual reproductive strategy is employed by this
species. It spawns in the piedmont of the Andes, during the low water
period of the dry season. Millions of tiny (2-8 mm), recently-hatched
larvae were collected by Alex Flecker and collaborators while sampling
aquatic insect drift at night, in the Las Marias and Masparro rivers
near Guanare during the peak of the dry season in January and February.
This phenomenon has been repeated now two years in row, and thus seems
to be a regular occurrence. This extremely unusual mode of breeding
has yet to be explained. The larvae are present at low water, when
predators would seem to be concentrated in the little remaining water
flow. I was able to rear a few larvae to a large enough size to permit
identification. To my knowledge, this is the only characiform fish
reported to have this mode of reproduction.

Figure 91. Apure drainage distribution of Bryconamericus deuterodonoides.


242
Importance. It has minor potential as an ornamental.
Bryconops (Creatochanes) affinis (Giinther) 1864
Sardinita - Tetra
Figs. 42b & 92. Map: fig. 93. Couplet 48b.
Generic synonymy
Bryconops Kner 1859:43 (type species: Bryconops alburnoides Kner
1859), by original designation.
Creatochanes Giinther 1864:329 (type species: Salmo melanurus (Bloch)
1794, by subsequent designation of Eigenmann 1910:435.
Specific synonymy
Tetraqonopterus affinis Giinther 1864:327 (type locality: British
Guiana), seen.
Tetraqonopterus rutilus (not of Jenyns) (part), Boulenger 1900:2.
Creatochanes affinis Eigenmann 1891:56; 1910:435; 1912:345; 1918:pi. 2,
fig. 5; 1921: pi. 23, fig. 3; 1922:237; Eigenmann & Myers 1929:436;
Norman 1926:92 (key); Tortonese 1941-42:117.
Bryconops (Creatochanes! mel anurus Müller & Troschel (not of Bloch)
1848:635.
Creatochanes melanurus Eigenmann & Eigenmann 1891:56; Eigenmann
1910:435; Steindachner (part) 1915:35.
Bryconops melanurus Bertoni 1939:55 (Paraguay).
Bryconops (Creatochanes! affinis Knóppel, Junk & Géry 1968:245 (key);
Géry 1977:435 (key).
Comments. The identification of this species is tentative. It
keys to B. affinis in Géry (1977), but doesn’t agree very well with the
original description, especially in relation to the lateral-line scale
count. It is probably a new species.

243
Figure 92. Brvconops (Creatochanes) affinis.

244
Etymology. BRYCON = is from Brycon a characid genus, OPS = like;
AFFINIS = similar to.
Description
Illustrations. Figs. 42b (maxilla length) & 92; Kndppel et al.
1968:233 (photo), 242 (caudal pattern); Géry 1977:435 (head).
Piagnosis. The species of Bryconops are long, spindle-shaped
tetras, with a long maxilla that extends well behind the anterior edge
of the orbit and well below the ventral edge of the orbit. B. affinis
has more lateral-line scales (54-60) than B. melanurus (43-45), the only
other species found so far in the Apure drainage. The pigment in the
caudal fin is more symmetrical than in B. melanurus, with both lobes
being darkest along their outer edges, with diffuse lighter yellow or
orange near each lobes’ base.
This species might be confused with Iquanodectes spilurus, but
lacks the characteristically shaped, multicuspid teeth of that species.
Size. This fish is usually taken at about 60 mm SL, but grows to
almost 90 mm.
Morphology. The dorsal-fin origin is behind the pelvic-fin
insertions. The lower caudal lobe is slightly longer than the upper.
Counts. DR ii9; AR iii31-iii33, (but given as 27 for type); LLS
54-60 (but given as 43 for type); PR i 11 - i 12; VR i8; TS 8-9 above, 4-4
1/2 below LL.
Measurements. HL 23-25% SL; GBD 23-25% SL.
Pigmentation. The body is yellowish in preservative with a wide
silvery lateral band from the opercle to the tail. The dorsal midline
is marked with a black line. The dorsal, anal, pectoral and pelvic fins
are clear, and often contain a few scattered melanophores along the
rays.

Figure 93. Apure drainage distribution of Bryconops affi


247
Distribution and Natural History
Range. It is reported from the Amazon, Orinoco, and coastal
drainages of Brazil, Venezuela, Guyana, French Guiana and Surinam.
Apure distribution. Map: fig. 93. It is known from only one site
in the Aguaro River system.
Habitat. This fish inhabits blackwater streams and morichales of
sandy savannas.
Abundance. RARE.
Number of specimens examined. 11 from 1 collection.
Food. OMNIVORE. It probably feeds on aquatic and terrestrial
insects that fall to the water’s surface or drift down with the current.
It is usually observed actively swimming about in large schools.
Reproduction. Strategy: probably rl. It probably spawns
repeatedly during the wet season.
Importance. Potentially valuable as an ornamental.
Bryconops (Creatochanes) melanurus (Bloch) 1794
Sardinita - Tetra
Fig. 94. Map: fig. 95. Couplet 88b.
Specific synonymy
Salmo melanurus Bloch 1794: tab. 381 fig. 2 (type locality: Surinam),
seen.
Tetragonopterus mel anurus Milller & Troschel 1845:14; Cuvier &
Valenciennes 1848:155.
Creatochanes melanurus Eigenmann 1910:435; 1912:346; Norman 1926:92
(key); Eigenmann & Myers 1929:438 (description).
Bryconops (Creatochanes) melanurus Kndppel, Junk & Géry 1968:245 (key);
Géry 1977:435 (range, key, description).

248
Figure 94. Brvconops (Creatochanes) melanurus.

249
Etymology. MELAN = black; URUS = tail.
Description
Illustrations. Fig. 94; Eigenmann 1912:pi. 50; Kndppel et al.
1968:232 (photo), 242 (caudal pattern); Géry 1977:420, 435 (head).
Diagnosis. The species of Bryconops are all long, spindle-shaped
tetras, with a long maxilla that extends well behind the anterior edge
of the orbit of the eye, and well below the ventral edge of the orbit.
In addition, B. melanurus has about ten fewer lateral-line scales (43-
45) than B. affinis (54-60), the only other species found so far in the
Apure drainage. The pigment in the caudal fin is concentrated in the
upper lobe.
This species might be confused with Iguanodectes spilurus but lacks
the characteristically shaped, multicuspid teeth of that species and
also has fewer lateral-line scales.
Size. It grows to about 60 mm SL.
Morphology. The greatest body depth measures 31/2-4 times in
the standard length. The lateral line is complete and decurved in
front. The origin of the dorsal fin is over the pelvic fin insertions.
Counts. DR ii9; AR i ii24-i ii26; PR i12-i13; VR i7; LLS 43-45; TS 7
above and 3 below LL.
Pigmentation. The body is yellowish in preservative, with a wide,
silvery black lateral band from the opercle to the tail. The dorsal-
midline is marked with a black line. The upper lobe of the caudal fin
is edged in black and has a yellow to orange spot near its base. The
lower caudal lobe is light colored, usually yellowish and the lateral
stripe continues onto the central caudal-fin rays. The other fins are
clear. The scales of the upper sides are outlined with gray.

Figure 95. Apure drainage distribution of Bryconops melanurus.

ro
en

252
Distribution and Natural History
Range. It occurs in the Amazon, Orinoco and coastal drainages of
Brazil, Venezuela, the Guianas and Surinam.
Apure distribution. Map: fig. 95. This fish is known only from
the Aguaro River system, the easternmost in the Apure drainage.
Habitat. This species frequents the blackwater streams and
morichales of the sandy savannas in the easternmost part of the Apure
drainage.
Abundance. UNCOMMON. This species is abundant where found.
Number of specimens examined. 256 from 19 collections.
Food. OMNIVORE. This fish probably feeds on aquatic and
terrestrial insects that fall to the water’s surface or drift down with
the current. It is usually observed actively swimming about in large
schools.
Reproduction. Strategy: probably rl. It probably spawns
repeatedly during the rainy season.
Importance. It is potentially valuable as an ornamental.
Catoprion mentó (Cuvier) 1819
Caribe Jetudo - Wimple Piranha
Fig. 96. Map: fig. 97. Couplet 20b.
Generic synonymy
Catoprion Müller & Troschel 1844:96 (type species: Serrasalmus mentó
Cuvier 1819, by monotypy).
Specific synonymy
Serrasalmus mentó Cuvier 1819:369 pi. 28 fig. 3 (type locality: Brazil?)
(not seen).
Catoprion mentó Müller & Troschel 1844:96 (British Guiana); 1845:22 pi.

253
Figure 96. Catoprion mentó.

254
2, figs 5, 5a; Eigenmann 1912:387, pi. 56, fig. 3 (key, description,
Essequibo); Mago L. 1970:70; Géry 1972:207 (diagnosis); 1977:294
(key).
Comments. This genus is monotypic, but I suspect that closer
examination of the different populations from the Amazon, Orinoco and
Guyana will reveal at least subspecific differences. I have noticed
considerable differences in lateral-line scale counts, number of ser-
rae, and other meristic character between the Apure population and
those reported from the Guianas (Géry 1972).
Etymology. CATO = probably from the Greek "kata" for lower,
PRION = is from the Greek for saw, perhaps referring to the saw-toothed
ventral keel in this species; MENTO = is from the Latin for chin,
calling attention to the jutting lower jaw, or chin of this species.
Description
Illustrations. Fig. 52; Eigenmann 1912:pi. 56, fig. 3; Axelrod
et al_. 1971:F-164.00; Myers 1972:64, 85, 124; Géry 1977:293, 294;
Bléher 1986:82, 84.
Diagnosis. C. mentó is distinguished by a prominent, protuberant
lower jaw and in having the first dorsal ray filamentous in larger,
undamaged specimens. It is further characterized by its unusual tuber-
cle-like teeth. The premaxilla has two large antrorse, conical teeth
in the "outer" row, and three much smaller teeth in the "inner" row,
which was called a "single irregular row" by Géry (1972). The mandible
only has a single series of teeth.
Size. This fish grows to about 100 mm SL.
Morphology. The dorsal fin and anal fin are falcate, and in some
aquarium specimens, a long filament forms on the dorsal fin. The body

255
is somewhat discoid and highly compressed. The belly is serrated as in
most piranhas.
Counts. DR ii14-16; AR 37-39; LLS 89-94; ventral serrae 32-34; GR
14.
Measurements. GBD 66% SL; Eye diameter about 2.7 in HL and equal to
interorbital width, longer than snout.
Pigmentation. The body is silvery, with a faint humeral spot and a
red spot on the opercle. The base of the caudal fin is black and the anal
fin is reddish, with its first rays white and elongated.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins, from southern
Brazil north to Venezuela, and in Guyana.
Apure distribution. Map: fig. 97. It is known only from northern
Apure state and the Guariquito system in Guárico state.
Habitat. In the Apure drainage, Catoprion mentó is restricted to
blackwater streams with abundant aquatic vegetation. To be able to feed
effectively, this species may need higher water clarity for enhanced
visibility.
Abundance. UNCOMMON.
Number of specimens examined. 187 from 24 collections.
Food. CARNIVORE. Lepidophage. This species is known to feed almost
exclusively on scales but it also may eat small fishes and insects (Nico &
Taphorn 1986, 1988; Vieira & Géry 1979).
Reproduction. Strategy: probably r2. This species is probably
an egg scatterer that spawns each year at the beginning of the rainy
season.
Importance. Ornamental.

Figure 97. Apure drainage distribution of Catoprion mentó.


Charax gibbosus (Linnaeus) 1758
Dientón, Jibao - Glass Headstander
258
Figs. 30 (head) & 98. Map: fig. 99. Couplet 7b.
Generic synonymy
Charax Scopoli 1777:455 (type species: Salmo gibbosus Linnaeus 1758, by
subsequent designation of Eigenmann 1910:444).
Characinus Lacepede 1803:269 (type species: Salmo gibbosus by subsequent
designation of Jordan & Evermann 1917:66).
Edicyrtus Müller & Troschel 1844:92 (type species: Salmo gibbosus
Linnaeus, by monotypy).
Anacyrtus Günther 1864:345 (type species: Salmo gibbosus Linnaeus, by
subsequent designation of Jordan 1919:333).
Specific synonymy
Salmo gibbosus Linnaeus 1758:311 (type locality: Surinam), seen.
Epicyrtus gibbosus Müller & Troschel 1844:92 (type locality: South
America).
Cynopotamus gibbosus Valenciennes, jn Cuvier & Valenciennes 1849:(240)
321.
Anacyrtus gibbosus Eigenmann & Eigenmann 1891:56.
Characinus gibbosus Eigenmann & Kennedy 1903:525.
Charax gibbosus Eigenmann & Ogle 1907:32; Eigenmann 1912:400 (synonymy,
key); Hensel 1962:127 (clarification of name); Géry 1977:301,307.
Roeboides microlepis Román 1985:161 (misidentified, Venez.).
Etymology. CHARAX = from the Greek for a palisade (Eigenmann
1927), in this case a palisade of teeth, but also given to mean a
"gnasher of teeth" in Eigenmann’s earlier works. Charax is a genus of
characid frequently used in the formation of generic names; GIBBOSUS =
refers to the hump-backed profile of the
nape.

259
Figure 98. Charax qibbosus.

260
Description
Illustrations. Figs. 30 (head) & 98; Eigenmann 1912:pi. 60, fig. 3;
Hensel 1962:128; Axelrod et al. 1971:F-171.00; Géry 1977:301. Román
1985:160, (misidentified as Roeboides microlepis. a species not known from
Venezuela).
Diagnosis. The characteristic humped back is the first clue to the
identity of this species. Small individuals of Roeboides. Cynopotamus and
Charax can be very difficult to distinguish. This species is distinguished
from Cynopotamus biounctatus by possessing cycloid rather than ctenoid
scales, and in having only 53-60 lateral-line scales versus over 100. It
may be distinguished from Roeboides spp. by the lack of everted teeth or
"tusks" on the upper jaw, and the presence of an elongated spiny projection
of the cleithrum below the base of the pectoral fin (which is triangular or
rounded in Roeboides spp).
Size. The glass headstander can grow to about 150 mm TL, but most
individuals are between 60-100 mm.
Morphology. The premaxilla has a feeble canine at either end and a
double row of conical teeth between, and the dentary has two feeble canines
and one series of conical teeth (Eigenmann 1912). The cleithrum has a
large notch to accommodate the pectoral-fin base. The dorsal-fin origin
is behind the anal-fin origin, but is nearer to the snout tip than to the
base of the caudal fin. There is no ventral keel, and the breast is flat.
The maxilla is longer than the eye diameter.
Counts. DR ii8-ii9; AR 43-56; VR 8; LLS 53-60; GR 8.
Measurements. GBD 32-43% SL; HL 23-24%; eye diameter equal to snout
length and to interorbital width in adult, 3.7 in HL;
Pigmentation. The body is whitish or silvery in preservative,
almost transparent in life, and the humeral spot is vertically elongate

Figure 99. Apure drainage distribution of Charax gibbosus.

no
en
IX)

263
and farther back on the side than in similar species (over the seventh
or eighth scale of the lateral line). The caudal fin has a faint
vertical bar at its base.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins, and the Guianas
(Géry 1977).
Apure distribution. Map: fig. 99. Charax qibbosus is widespread in
the lower parts of the drainage.
Habitat. It is usually found in the sluggish areas of lowland
creeks, swamps and other lentic habitats.
Abundance. COMMON.
Number of specimens examined. 984 from 144 collections.
Food. CARNIVORE. This species eats small fish and aquatic
insects, especially Hemiptera (K. Winemiller pers. com.).
Reproduction. Strategy: r2; generation time: 6 months; fecundity:
280 eggs/female; egg diameter: 1.00 mm (Winemiller & Taphorn 1989). Eggs
are scattered over aquatic vegetation and then abandoned. Incubation
lasts 30 hours, and after the yolk sac is absorbed the fry feed on plank¬
ton (Axelrod et al. 1971).
Importance. Ornamental.
Cheirodon pulcher (Gill) 1858
Sardinita - Pretty Tetra
Figs. 53 (teeth) & 100. Map: fig. 101. Couplet 77a.
Generic synonymy
Cheirodon Girard 1854:199 (type species: Cheirodon pisciculus Girard
1854, by monotypy).

264
Figure 100. Cheirodon pulcher.

265
Chirodon Giinther 1864:332 (type species: Cheirodon pisciculus Girard
1854), (alternative spelling).
Odontosti 1 be Cope 1870:566 (type species: Odontosti 1 be fugitiva Cope
1870, by monotypy).
Specific synonymy
Poecilurichthvs pulcher Gill 1858:422 (type locality: Trinidad), seen.
Chirodon (Odontostilbe) pulcher Liitken 1874:238 (Venezuela?);
Odontosti 1 be pul cher Eigenmann 1920:9 (Maracay area, Venez.); Schultz
1944b:318 (key, Caripito, Venez.).
Odontostilbe pulchra Géry 1977:558 (key; taxonomic comments).
Comments. The question of the proper name for this genus is still
unresolved. Fink & Weitzman (1974) synonymized Odontosti1 be with
Cheirodon. but Géry (1977) maintained the genus Odontostilbe for those
forms with a complete lateral line.
Etymology. CHEIR = hand shaped, ODON = teeth. PULCHER = pretty.
Description
Illustrations. Fig. 53 (teeth) & 100; Regan 1906:pi. 22.
Diagnosis. This species can be distinguished from similar small
tetras (for example Hemigrammus) by having its premaxillary teeth in a
single row. The lateral line is incomplete. The very similar
Cheirodontops geayi is more slender and has a smaller caudal spot, a
complete lateral line, and differs markedly in dentition. Cheirodon
pulcher has premaxillary teeth with nine cusps, and dentary teeth with
seven. The teeth in the upper and lower jaw are essentially similar in
shape, with the central cusp longest. Cheirodontops geayi, on the other
hand, usually has premaxillary teeth with only five cusps, and very
different dentary teeth (three flattened central cusps, and two tiny
outer cusps on each tooth) see fig. 53.

266
Size. This is a small fish that seldom exceeds 30 mm SL.
Morphology. The dorsal fin is equidistant from the snout and the
base of the tail. The maxilla extends back to the level of the front
margin of the eye. There is a short sheath of scales on the anterior
base of the anal fin.
Counts. DR ii9; AR iii20-iii22; PR 13; VR i7; LS 32-35; TS 11 (6
above 5 below LL). Premaxilla with 5-6 teeth, maxilla with three,
dentary with seven, seven cusps on each tooth in both jaws, sometimes
the teeth have black or reddish tips.
Measurements. Body depth 2.5-2.8 in SL; HL 3.7-4.7 in SL.
Pigmentation. There is distinct black spot at the base of the
caudal fin that extends onto the caudal peduncle. A red, yellow or
orange dot is present above (and often below as well) this. The eye is
red above the pupil and there is a gold spot on and behind the opercle.
A narrow lateral stripe starts at or just in front of the dorsal-fin
origin. All the scales above this line are specked with pigment
(concentrated posteriorly on each scale), but those below are nearly
white.
Distribution and Natural History
Range. This species is found from Trinidad, and the Orinoco Basin
of Venezuela and Colombia.
Apure distribution. Map: fig. 101. This small tetra is ubiquitous
in the piedmont and llanos.
Habitat. It occurs in almost any water body, but is more common in
lowland lentic systems.
Abundance. ABUNDANT. This is the most ubiquitous species in the
Apure drainage.

Figure 101. Apure drainage distribution of Cheirodon pulcher.

no
cr>
co

269
Number of specimens examined. 21,330 from 431 collections.
Food. OMNIVORE. The diet of this species includes filamentous
algae, microcrustaceans, protozoa, desmids and diatoms (K. Winemiller,
pers. com.).
Reproduction. Strategy: rl; generation time: four months;
fecundity: 1108 eggs/female; egg diameter: 0.75 (Winemiller & Taphorn
1989). These fish spawn throughout the rainy season, and individuals
may spawn more than once. It is an egg scatterer.
Importance. Ornamental.
Cheirodontops qeavi Schultz 1944
Sardinita - Tetra
Figs. 53 (teeth) & 102. Map: fig. 103. Couplet 77b.
Generic synonymy
Cheirodontops Schultz 1944b:319 (type species: Cheirodontops geayi
Schultz 1944b, by original designation).
Specific synonymy
Cheirodontops qeayi Schultz 1944b:319 fig. 42,43 (type Locality: Rio
Guárico and tributaries between San Sebastian and San Casimiro,
Aragua, Venez.), seen; Mago L. 1970:70; Géry 1977:546 (key).
Types. Holotype: USNM 121507. Paratypes: CAS-SU 18144 (formerly
USNM 121508).
Comments. As Schultz himself noted (1944b), the only difference
between the genera Cheirodontops Schultz 1944b and Holoshesthes Eigen-
mann 1903 is the presence of one or two maxillary teeth in the latter.
Schultz examined only seven specimens to describe the new genus. I
doubt that this small difference warranted the description of a new
genus.

270
Figure 102. Cheirodontops qeayi.

271
Etymology. CHEIRODON = Cheirodon, a related and similar genus of
cheirodontine characid, OPS = like; GEAYI = after French biologist F.
Geay, who reported on the fisheries of the Orinoco Basin in his work
"Peches dans les Affluentes de 1’Orinoque" (Schultz 1944b).
Description
111ustrations. Figs. 53 (teeth) & 102; Schultz 1944b:320, (this
figure shows the fish as much deeper bodied and stockier than most
specimens I have seen).
Di agnosis. This species can be distinguished from many similar
small tetras (for example Hemigrammus spp.) present in the Apure
drainage by the presence of a single row of premaxillary teeth. It is
similar in this respect and in overall appearance to Cheirodon pulcher,
but can be distinguished by its complete lateral line, more slender
body, smaller caudal spot and in particular, the very different
dentition. Cheirodontoos geavi have differently shaped teeth on the
premaxilla and dentary. While most teeth of both jaws are pentacuspid,
the outer cusps are often minute, those on the dentary have three, wide,
flat central cusps, whereas those on the premaxilla have the central
cusp pointed, and larger than the others. See Schultz (1944b) for a more
detailed description.
Size. This fish can grow to about 50 mm SL, but most individuals
are between 30-40 mm.
Morphology. The anal-fin base has a sheath of scales anteriorly.
Counts. DR ii-9; AR ÍÜ21-22, hooks present on rays in males; PR
11; VR i7; branched caudal-fin rays 17; LLS 35-37; TS 10-11, five above
and four below the lateral line. Premaxillary teeth six on either side,
dentary with 4 large teeth and six smaller on each side; GR 14.

272
Measurements. HL 4.1-4.3 in SL; GBD 3.0-3.4 in SL; EYE 3.0-3.2 in
HL.
Pigmentation. The body is silvery to whitish. On the dorsum there
are three rows of scales which have white centers outlined in gray. The
fins have very little pigment. The caudal spot extends from the base of
the central caudal-fin rays onto the caudal peduncle. The lateral
stripe consists of both deep (embedded) and superficial pigment and
there is also a concentration of pigment along the base of the anal fin.
Distribution and Natural History
Range. So far this species is reported only from Venezuela, in the
Orinoco basin but it is undoubtedly present in the Colombian part of the
basin as well.
Apure distribution. Map: fig. 103. This fish is widely
distributed throughout the lower portion of the drainage.
Habitat. This fish seems to be at home in main river channels,
smaller streams, and even in standing waters.
Abundance. COMMON.
Number of specimens examined. 1,705 from 101 collections.
Food. CARNIVORE. This species feeds on microcrustaceans and
aquatic insects.
Reproduction. Strategy: r2; generation time: 12 months; fecundity
1108 eggs/female; egg diameter 0.75 mm (Winemiller & Taphorn 1989).
This species spawns annually at the onset of the rains. It is probably
an egg scatterer with no parental care.
Importance. Ornamental, but not highly sought due to the absence
of "interesting" colors.

Figure 103. Apure drainage distribution of Cheirodontops geayi.


275
Colossoma macropomum (Cuvier) 1818
Cachama - Pacu
Fig. 39a (operculum) & 104. Map: None. Couplet 30a.
Generic synonymy
Colossoma Eigenmann & Kennedy 1903:350 (type species: Mvletes oculus
Cope 1872, by original designation).
Waiteina Fowler 1907:473 (type species: Mvletes nigripinnis Cope 1878 by
original designation).
Regañina Fowler 1907:475 (type species: Myletes bidens Spix i_n Agassiz
1829, by original designation).
Colosoma Eigenmann 1917:35. (misspelling).
Mellonina Campos 1946:219 (type species: Mellonina tambaaui Campos 1946,
by original designation).
Specific synonymy
Mvletes macropomus Cuvier 1818:453, pi. 22 fig. 3 (type locality:
Brazil), seen; Valenciennes in Cuvier & Valenciennes 1849:(146)195
(redescription of type); Peters 1877:473 (Apure River, Venez.);
Eigenmann & Eigenmann 1891:61.
Myletes bidens (not of Spix) Castelnau 1855:68 pi. 35 fig 1-3.
Salmo tambaaui Natterer jn Kner 1860:21 (mentioned in text).
Myletes oculus Cope 1872:268, pi. 12 fig. 3 (type locality: Ampiyaco,
Peru).
Myletes nigripinnis Cope 1878:693 (type locality: Peru, Pebas, Nauta);
Steindachner 1882:125 pi. 7, fig. 1 (redescription).
Colossoma oculus Eigenmann & Kennedy 1903:530 (new generic name for
Myletes oculus Cope 1872); Fowler 1906:472 (redescription); Norman
1929:811 (redescription, synonymy, Peru).

276
Figure 104. Colossoma macropomum.

277
Colossoma (Waiteina) nigripinnis Fowler 1906:473.
Colossoma nigripinnis Norman 1929:808 (key, synonymy).
Piaractus macropomus Eigenmann 1910:443 (R. Orinoco); Román 1985:113
(description, Venez.).
Piaractus nigripinnis Eigenmann 1915a:262 (redescription); Eigenmann &
Allen 1942:247.
Colossoma macropomus Norman 1929:808 (key, synonymy); Schultz
1944b:255; Fowler 1950; Mago L. 1970:70.
Mellonina tambaoui Campos 1946:219 fig. 3 (type locality: ?Brazil).
Col ossoma (Piaractus) brachypomum Géry 1977:255.
Colossoma macropomum Machado A. 1982: (diagnosis, description,
synonymy).
Types. Holotype: MNHN A.8627.
Etvmology. C0L0S = without horns (that is, lacking a predorsal
spine, as is present in many serrasalmines), SOMA = body; MACRO = large,
P0MUM = cheek, referring to the large, wide operculum.
Description
Illustrations. Fig. 39a (operculum) & 104; Géry 1977:255; Novoa et
al_. 1982: 269, fig. 6; Machado A. 1982: (photos of holotype and
juveniles); Román 1985:113.
Diagnosis. There are only two species of cachamas known in the
Apure drainage. C. macropomum is distinguished from Piaractus
brachypomus in having a wider operculum (see fig. 39a), fewer lateral¬
line scales (67-76 vs 78-90), fewer scales between the dorsal-fin origin
and the lateral line (19-23 vs 26-30) and by the presence of ossified
rays in the adipose fin of specimens larger than 55 mm SL.
Size. Specimens measuring 920 mm TL and weighing 18 kg have been
reported from the Orinoco Delta (Novoa et al. 1982).

278
Morphology. This is a giant characid, with a heavy set, discoid
body. See Machado A. (1982) for a detailed account.
Counts. DR ii-ivl3-14 (15-18); AR i i-iv21-24 (24-28); PR i15-18
(16-19); VR i7; caudal fin i-9+8-i; LLS 66-78; TS 16-23 above LL, 19-27
below LL; GR vary with size from about 20 in a specimen of 19 mm SL to
136 in a large adult of 755 mm SL (Machado A. 1982).
Measurements. GBD 45.0-67.6% SL; HL 34.4-51.9% SL (Machado A.
1982).
Pigmentation. Juveniles up to 30 mm SL have a series of irregular
spots over a yellowish background, with a large ocellus on the rear half
of the flank, and a reddish head. The caudal-fin is edged in black and
the anal fin is orange with a black border. The other fins are mostly
transparent. As the fish grow, the base color darkens and the ocellus
and irregular spots are lost. The fins eventually darken to black.
Adults have the ventral region dark or black, the lateral and dorsal
regions olivaceous or copper-colored, and the fins black (Machado A.
1982).
Goulding (1980) noted that this species is countershaded in
Amazonian waters, but backwards relative to most fishes. It is lighter
above and darker below. The colors of specimens taken from blackwater
streams are more intense.
Novoa et al. (1982) reported two different color morphs present in
the Orinoco Delta area. In one the dorsum is yellow and the ventrum
white; in the other, the fish is mostly black with a whitish belly. I
have observed more dark-colored fish in the Apure markets, with a
generally black coloration that is most intense on the pectoral fins,
lower sides, anal fins, lower caudal peduncle and tail. The belly is
wh i t i s h .

279
Distribution and Natural History
Range. It occurs in the Orinoco and Amazon basins.
Apure distribution. Map: None. No specimens were obtained during
this study, mainly because commercial seining gear or hook and line are
usually necessary to catch them. It is (or was until recently) fairly
common in most of the larger rivers.
Habitat. In the Apure drainage, this species is caught mostly in
the main channels of large rivers during migrations, and in larger
llaneran creeks during the dry season. The cachama fishery is
endangered because of the loss of habitat due to increased drainage of
flooded forests, dikes that prevent access to the floodplain, the
continuous loss of the gallery forests that once provided the fruits and
seeds essential to its survival, and the overfishing of ripe females
during the dry season.
Abundance. COMMON.
Food. HERBIVORE. This species eats large seeds, fruit, flowers,
aquatic vegetation. Goulding (1980) noted that their fine gillrakers
are used to trap plankton when young. Adults use massive dentition to
crush large heavy seeds, such as those of the rubber tree Hevea and the
palm Astrocarvum. These trees are widely distributed in the Amazon
basin and produce large, numerous, high protein seeds which are too big
or hard for most other species to eat. The cachamas actively seek out
trees in fruit. When food becomes scarce during low water, their diet
increases in variety of foods eaten. Thus, a dietary specialization is
in effect mainly during the "good" period of the year when food is
abundant, not during the times of scarcity as some authors have
suggested.

280
Reproduction. Strategy: r2. Gonads ripen during the dry season
and mating is synchronous with the onset of the rains in May to June.
Novoa et al. (1982) reported that cachamas in the Orinoco Delta area
move onto the inundated floodplains to spawn in the shallows. Fry stay
in the aquatic vegetation where they grow quickly, feeding on plankton
and algae. At the end of their first wet season, young fishes move from
the floodplain back into regular watercourses. Fecundity is very high;
the largest females can produce around one million eggs each year.
Migrations. Extensive migrations are known to occur, but different
regions have different movement patterns. In the Apure drainage, fish
tend to move upstream in the dry season, and back down with the rains.
Importance. This is an extremely important species in the
commercial fishery, and of growing importance in fish culture. It is
usually captured with large meshed seines operated from two dugout
canoes with outboard motors or (much less frequently) with gill nets.
Corvnopoma riisei Gill 1858
Sardinita Cachete de Espada - Sword-Cheeked Tetra
Fig. 105. Map: fig. 106. Couplet 2a.
Generic synonymy
Corvnopoma Gill 1858:65 (type species: Corvnopoma riisei Gill 1858, by
original designation); Regan 1906:382.
Stewardia Gill 1858:63 (type species: Stewardia albipinnis Gill 1858,
by original designation); Eigenmann 1914:37.
Nematopoma Gill 1858:68 (type species: Nematopoma searlesii Gill 1858,
by original designation).
Specific synonymy
Corvnopoma riisei Gill 1858:66 (type locality: Trinidad), seen;

281
Figure 105. Corvnopoma riisei.

282
Eigenmann & Myers 1929:470 (key, description); Regan 1906:382
(redescription); Axelrod et al. 1971:F-225.00; Mago L. 1970:70; Géry
1977:356 (key); Richter 1986:44 (spawning).
Stewardia albipinnis Gill 1858:65 (type locality: Trinidad).
Corynopoma albipinne Giinther 1864:287.
Corynopoma veedonii Gill 1858:67 (type locality: Trinidad).
Nematopoma searlesii Gill 1858:69 (type locality: Trinidad).
Corynopoma searlesii Giinther 1864:288; Liitken 1874:222 (Trinidad).
Stewardia aliata Eigenmann 1914:37 (type locality: base of Andes east of
Bogotá); 1922:238.
Corynopoma aliata Eigenmann & Myers 1929:468 (key, description).
Etymology. C0RYN0 = spatula, POMA = cheek or operculum, meaning
"with a spatula attached to the gill plate"; RIISEI = after Mr. A. H.
Riise, a Danish zoologist who worked in the West Indies.
Description
Illustrations. Fig. 105; Regan 1906:pi. 22; Axelrod et al.
1971:F-225.00; Géry 1977:356, 357; Mago L. 1978:9; Richter 1986:44-5,
48-9.
Diagnosis. This fish strongly resembles Gephyrocharax valenciae,
with which it is frequently sympatric, but differs in lacking an adipose
fin. The strange filamentous extension of the opercle in males of this
species is unique among Venezuelan characids, although similar
structures have been described for Pterobrycon, a Central American
glandulocaudine characid (Bussing 1971).
Size. Males grow to about 75 mm TL, but females are smaller.
Morphology. The mouth is superior, with the lower jaw extending
beyond the upper. The nostrils are near the antero-dorsal corner of

283
eye. The lateral line is initially deflected downward after which it
continues straight to the base of the caudal fin. The dorsal fin is
placed well back, with its origin over the middle of the anal-fin base.
The caudal fin is deeply forked.
Counts. DR 8; AR 27; PR 10; VR 6.
Pigmentation. This is a silver-bodied to pinkish fish with clear
fins. There is a faint horizontal stripe that is most intense and widest
on the caudal peduncle. The tip of the opercular "paddle" is shiny.
Distribution and Natural History
Range. It occurs in Trinidad, and in the Venezuelan and Colombian
Andes.
Apure distribution. Map; fig. 106. This species is strictly
limited to the Andean piedmont.
Habitat. This fish inhabits shady, shallow, slow-moving streams of
the Andean piedmont.
Abundance. COMMON.
Number of specimens examined. 362 from 28 collections.
Food. CARNIVORE. It feeds mostly on terrestrial insects,
especially ants (K. Winemiller pers. com.)
Reproduction. Strategy: rl; generation time: 4 months; fecundity:
135 eggs/female; egg diameter 0.85 mm (Winemiller & Taphorn 1989).
Individuals spawn repeatedly throughout most of the year.
The courtship ritual is elaborate, with much display activity by
the male. Fertilization is internal, but the mechanics by which this is
effected is as yet unexplained. The male wraps his anal fin around the
female to form a pocket as he expels the sperm, but the female does not
expel the eggs at that time. She will later place the embryos in

Figure 106. Apure drainage distribution of Corynopoma riisei

285

286
aquatic vegetation. They hatch in about 30 hours, and are free-swimming
after about 4 days, when they will start to feed on plankton (Richter
1986). Dr. Stanley Weitzman (pers. com.) has studied the reproductive
habits of this and other glandulocaudines. He believes that the "tail
beat" movements of the courtship dance may direct the pheromone produced
in the caudal glands towards the female and that the enlarged, modified
scales on the caudal fin base of males may act as pumps for the
pheromone. During the nuptial dance, the female nips at the opercular
"paddles." This may polarize her attention sufficiently for the male to
fertilize her.
Importance. Ornamental.
Creaqrutus cf beni Eigenmann 1911
Dientefrio - Buck-Toothed Tetra
Figs. 40 (humeral spot) & 107. Map: fig. 108. Couplet 37b.
Generic synonymy
Creaqrutus Günther 1864:339 (type species: Leporinus muelleri Giinther
1859:90, by monotypy).
Specific synonymy
Creaqrutus beni Eigenmann 1911:172, pi. 6, fig. 2 (type locality: Villa
Bella, Bolivia, Rio Beni into Rio Madeira [a southern tributary of the
Amazon River]), seen; Eigenmann 1918:pi. 93, figs. 4, 5, 7; 1920:12
(Maracay, Valencia and Caracas, Venez.); 1927:421 (Valencia, Venez.);
Pearse 1920:20,43 (Valencia, Venez.); Fowler 1931:408 (Yarapa River,
Venez.); 1940:100; 1942:128; 1945a:148; 1945b:102; 1948:83 (synonymy,
range); Schultz 1944b: (Maracaibo Basin, Rio Tuy in Caracas, and
Caripito, Venez.); Mago L. 1970:70; Géry 1977:392, 407, 410.
Types. Holotype: FMNH 54585, (formerly CM 3216), examined.

Figure 107. Creaqrutus cf beni.

288
Comments. The holotype of C. beni, FMNH 54585, from Bolivia, is
in poor condition based on my examination. Eigenmann’s photo (1911) of
the holotype shows a vertically elongated humeral bar, very different
from the concentrated spot or blotch observed in Venezuelan specimens
from the Apure drainage. I am convinced that the Venezuelan form
represents a different species. A Bolivian specimen, FMNH 77395, has a
crescent-shaped humeral spot, something almost never seen in Venezuelan
"C. beni."
Etymology. CREA = flesh, from the Greek, GRUTUS = to tear, also
Greek; BENI = for the river in Bolivia of the same name, type locality
for this species.
Description
Illustrations. Figs. 40 (humeral spot) & 107; Eigenmann 1911:pi.
6, fig.2; 1918:pi. 93 fig. 7 (photo).
Diagnosis. Members of the genus Creagrutus can be distinguished
from most other tetras in the Apure drainage by their prominent,
projecting upper jaws, and in possessing three (versus one or two) rows
of teeth on the premaxilla. Creagrutus cf beni has a rounded humeral
spot instead of a crescent-shaped one, as observed in C¿. bol i vari. It
is distinguished from Creagrutus n. sp. in having fewer scales between
the lateral line and the dorsal-fin origin (five vs six), and in usually
lacking the dark black pigment seen in the central area of the dorsal
fin of that species.
Size. This fish can grow to about 60 mm SL, but most individuals
are between 30-50 mm SL.
Morphology. The dorsal-fin origin is behind or over the pelvic-fin
origin. The lateral line is complete.

289
Counts. DR 10; AR i ii10-i ii12; LLS 35-40.
Measurements. HL 31/3-4 in SL; GBD 3-3 1/2.
Pigmentation. The humeral spot is rounded and not shaped like an
open vertically elongate crescent (although the rounded spot sometimes
can take the form of a tight, closed crescent). Usually there is a
concentration of pigment near the lateral line if vertical extensions of
the humeral spot are present. The body is pinkish in life, with a dark
lateral stripe. The dorsum does not have a complete line of darker
scales down the midline. There is no spot on the caudal peduncle.
Distribution and Natural Hi story
Range. Although this species is reported from the Andean
headwaters of the southern Amazon basin in Bolivia, northward to the
Maracaibo basin and the Orinoco basin in Venezuela, I suspect that
several similar species are involved.
Apure distribution. Map: fig. 108. This fish is restricted to
streams and rivers in the Andean mountains and piedmont.
Habitat. C. beni is found in piedmont and montane streams
throughout the Andes, in swiftly flowing waters over rocky substrates.
Abundance. COMMON.
Number of specimens examined. 1,315 from 87 collections.
Food. Benthic OMNIVORE. It feeds mostly on aquatic insects, and
small seeds (K. Winemiller pers. com.), but Pearse (1920) reported a
diet of 96.2% (volume) snails and 3.8% midge pupae from Lake Valencia.
I have observed them feeding in the pools and quieter stretches of
rapids where they usually are observed near the bottom in small groups
feeding on detritus, items rolling down in the current, and aquatic
insects. C. beni is extremely active, and always on the go.

Figure 108. Apure drainage distribution of Creagrutus cf beni.

no

292
Reproduction. Strategy: probably rl. Hoedeman (1975) stated that
fertilization is internal, but I have not observed any specialized
structures in males that might be uses to accomplish this. In aquaria,
eggs are laid among fine-leaved plants and hatch in about one day at
25°C. After three days in the yolk-sac stage, the fry begin to feed on
piankton.
Importance. Ornamental.
Creaqrutus bol i vari Schultz 1944
Dientefrio - Buck-Toothed Tetra
Figs. 40b & 109. Map: fig. 110. Couplet 37a.
Specific synonymy
Creaqrutus bol i vari Schultz 1944b:334, fig. 49 (type locality: Rio
Guárico and tributaries between San Sebastian and San Casimiro, State
of Aragua, Venezuela), seen; Myers & Roberts 1967:248; Mago L.
1970:70; Géry 1977:403,406 (key).
Comments. Based on observations by Myers and Roberts (1967), Mago
L. (1970) stated that this species is probably a junior synonym of C.
phasma. It is also similar to C. melanzonus Eigenmann 1912, (I examined
the holotype FMNH 52705), but that has a small caudal spot. I believe
that the species of Creaqrutus species will eventually prove to be
fairly restricted in distribution once a thorough revision of the genus
is completed. Because the habitats are so different, a more detailed
comparison of material from Andean streams with that from the Aguaro
River system should be undertaken in order to establish if these two
populations represent but a single species.
Etymology. BOLIVARI = after Simon Bolivar, liberator of northern
South America.

293
Figure 109. Creaqrutus bol i vari

294
Description
II1 ustrations. Figs. 40b (humeral crescent) & 109; Schultz
1944b:335 fig. 49.
Diagnosis, see previous account of C. cf beni.
Size. This fish grows to about 60 mm SL.
Morphology. The dorsal-fin origin is over or in front of the
pelvic-fin insertion. The lateral line is complete.
Counts. DR ii8; AR ii-iii8, ii-iiilO, but usually i ii9; PR i 11 -
i14; VR i7; LLS 37-38; TS 5 above LL, 3 below; PDS 9.
Measurements. HL 3 2/3 - 4 in SL; GBD 3 1/3 - 4; width of second
suborbital about 2/3 of eye diameter.
Pigmentation. The humeral spot is vertically elongate and cres¬
cent-shaped. The lateral stripe does not continue onto the caudal fin,
and there is no caudal spot. The dorsum has a few rows of dark spots
along the dorsal midline that contrast with the whitish upper sides.
The paired fins are plain.
Distribution and Natural History
Range. Orinoco Basin.
Apure distribution. Map: fig. 110. This species is widespread.
Habitat. C. bol i vari occurs in flowing waters of the piedmont and
llanos, usually over rocky substrates.
Abundance. COMMON.
Number of specimens examined. 759 from 71 collections.
Food. OMNIVORE. L_ bol i vari feeds on small benthic organisms and
those that drift down with the current.
Reproduction. Strategy: probably rl. Breeding probably occurs
throughout the rainy season.
Importance. Ornamental.

Figure 110. Apure drainage distribution of Creagrutus bol i vari.


Creaqrutus n. sp.
Dientefrio Pintada - Red-Finned Buck-Toothed Tetra
Fig. 111. Map: fig. 112. Couplet 36a.
297
Comments. This is probably a new species.
Description
II1ustrations. Fig. Ill; Géry 1977:392 (bottom photo); Román
1985:170.
Diagnosis. See account of C. cf beni. In addition, this species
has distinctive colors in life. The dorsal fin and anal fin have
patches of cherry-red anteriorly, and black pigment centrally. The
upper lobe of the caudal fin is also red.
Size. This is a relatively small Creaqrutus that can grow to about
50 mm SL, but most individuals are between 25-35 mm.
Morphology. The dorsal-fin origin is over the pelvic-fin
insertions. The lateral line is complete.
Counts. DR 10; AR i ii10-11; LLS 35; PDS 11.
Measurements. Body depth 2.9-3.3 in SL.
Pigmentation. In addition to the characters mentioned in the
diagnosis, this species has a lateral stripe that begins under the
dorsal fin and intensifies posteriorly. The humeral spot is vertically
elongate. The belly is whitish.
Pistribution and Natural History
Range. It probably occurs in both Venezuela and Colombia in the
Andean piedmont of the Orinoco Basin.
Apure distribution. Map: fig. 112. This species is restricted to
piedmont and montane stream.
Habitat. It lives in small, shallow, shady streams in the Andean
piedmont that usually have clear water and a sandy or gravel substrate.

298
Figure 111. Creagrutus n. sp.

Figure 112. Apure drainage distribution of Creagrutus n. sp.


301
Abundance. COMMON.
Number of specimens examined. 575 from 33 collections.
Food. OMNIVORE. This species feeds on tiny seeds and aquatic
insects (especially chironomid larvae) (K. Winemiller, pers. com.).
Reproduction. Strategy: rl; generation time: four months;
fecundity: 94 eggs/female; egg diameter: 0.80 (Winemiller & Taphorn
1989). It spawns throughout the wet season. Individuals probably spawn
more than once a season.
Importance. This strikingly colored species has great potential as
an ornamental fish. It is quite hardy in the aquarium.
Ctenobrvcon spilurus (Valenciennes) 1849
Sardinita Plateada - Silver Tetra
Fig. 113. Map: fig. 114. Couplets 10a & 40a.
Generic synonymy
Ctenobrvcon Eigenmann 1908:94 (type species: Tetragonopterus
hauxwellianus Cope 1870:560, by original designation).
Apodastvanax Fowler 1911:422 (type species: Apodastvanax stewardsoni
Fowler 1911:422, by original designation).
Specific synonymy
Tetragonopterus spilurus Valenciennes fn Cuvier & Valenciennes 1849:156
(type locality: Surinam).
Apodastvanax stewardsoni Fowler 1911:422 (type locality: Corisal,
Venez.).
Ctenobrvcon spilurus Eigenmann 1910:435; 1912:363; 1927:335 (Surinam to
Venezuela); Mago L. 1970:70; Géry 1977:413,431 (key).
Etymology. CTEN0 = rough scaled, BRYCON = Brycon, a genus of
characid; SPIL= spot, URUS = tail.

302
Figure 113. Ctenobrvcon spi 1urus.

303
Description
Illustrations. Fig. 113; Eigenmann 1912:pi. 47, fig. 1; Axelrod
et al. 1971:F-233.00; Géry 1977:413.
Diagnosis. This species is distinguished by the following combi¬
nation of characters: ctenoid scales, especially obvious on the breast;
a naked caudal-fin, usually with a black spot near its base; a double
row of teeth on the premaxilla, a complete lateral line, and a rela¬
tively high anal-fin ray count (about i ii37).
Size. It can grow to about 75 mm, but most individuals are
smaller.
Morphology. The swim bladder is two-chambered, with the anterior
shorter and rounded.
Counts. DR Ü9; AR iv37; PR i 12; VR i7; LLS 45; TS 21, 10 above
and 10 below lateral line from dorsal origin to pelvic origin; GR 12-14.
Measurements. GBD 3.2-3.5 in SL; HL 3.8-4.0 in SL.
Pigmentation. This fish is plain silvery to whitish with an
olive-green or sometimes golden sheen. The humeral and caudal spots
are variable, but usually the caudal spot is stronger. The fins are
transparent. The peritoneum is silver towards the ventrum, but pig¬
mented dorsally.
Distribution and Natural History
Range. It occurs in Surinam, Guyana and Venezuela (in the Orinoco
Basin).
Apure distribution. Map: fig. 114. This species is widespread and
common throughout the piedmont and llanos.
Habitat. C. soilurus favors lentic habitats such as prestamos,
bajios and esteros, but is also found in slow-moving streams.
Abundance. ABUNDANT.

Figure 114. Apure drainage distribution of Ctenobrycon spilurus.

305

306
Number of specimens examined. 16,691 from 355 collections.
Food. OMNIVORE. It eats seeds and aquatic insects.
Reproduction. Strategy: rl; generation time: 11 months;
fecundity: on the average about 755 eggs/female but up to 2,000; egg
diameter: 0.75 mm (Winemiller & Taphorn 1989). Spawning occurs during
the first two months of the rainy season. Individuals may spawn more
than once per season. Eggs are scattered among vegetation, and then
abandoned. Fry hatch within 50-70 hours, remain inactive while they
absorb the yolk-sac for 2-3 more days and then begin to feed on
pi ankton.
Importance. It is quite peaceful and hardy in aquaria.
Cvnopotamus bipunctatus Pellegrin 1909
Dientón
Fig. 115. Map: fig. 116. Couplet 41a.
Generic synonymy
Cvnopotamus Valenciennes 1849:316 (type species: Hvdrocvon argenteum
Valenciennes 1849), by original designation; Eigenmann 1912:403.
Cvrtocharax Fowler 1907:454 (type species: Anacvrtus 1 imaesouamis, by
original designation); Cope 1878:686.
Hybocharax Géry & Vu-Tan-Tue 1963b:240 (type species: Cvnopotamus
bipunctatus Pellegrin 1909:13, by original designation).
Specific synonymy
Cvnopotamus bipunctatus Pellegrin 1909:13 (type locality: mouth of the
Río Suripá, Venez.); Géry & Vu-Tan-Tue 1963a:150; Menézes 1976:31
(redescription of unique holotype, key).
?Anacvrtus humeralis Pellegrin 1899 (Apure River, Venez.), (this may
have been based on specimens of Galeocharax).

Figure 115. Cynopotamus bipunctatus. From Menézes (1976).

308
Cvnopotamus humeral is Schultz 1944b:296.
Cvrtocharax bipunctatus Schultz 1950:62.
Cvnopotamus (Hvbocharaxl bipunctatus Géry & Vu-Tan-Tue 1963b:241 (key);
Mago L. 1970:70.
Etymoloqy. CYNO = wolf, referring to the large canine teeth
characteristic of this species, POTAMUS = of the river; BI = two;
PUNCTATUS = spotted.
Description
Illustrations. Fig. 115; Menézes 1976:fig. 27.
Diagnosis. The absence of everted teeth that form "tusks" will
distinguish this species from Roeboides spp. It has a much more
pronounced humpback, and much smaller scales (over 100 in the lateral¬
line series vs fewer than 85) than the Acestrocephalus or Galeocharax
present in the Apure drainage.
Size. This species reaches at least 120 mm SL.
Morpholoqy. The pectoral-fin base shield is not particularly
developed, but the cleithrum is notched to receive the pectoral-fin
base. The dorsal-fin origin is in front of anal-fin origin. The scales
are ctenoid and the lateral line is complete.
Counts. DR i i9; AR Ü47-Ü49; PR i 14; VR i7; LLS 104-110; TS 26
above LL, 25 below; maxilla with 46 teeth; one row of large teeth on
mandible, the second "row" represented by only 2-3 conical teeth near
the symphysis. Anal sheath with 5 rows anteriorly, but reduced to 2
posteriorly.
Pigmentation. The humeral spot is rounded to oval, located near
the opercle, and is larger than the eye. The caudal spot is small and
rounded. The lateral stripe is silvery, narrow at both ends, and wider
in the middle. The fins are clear.

Figure 116. Apure drainage distribution of Cynopotamus bipunctatus.

o

311
Distribution and Natural Hi story
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 116. This is a low piedmont and
llanos species with a spotty distribution.
Habitat. It is usually taken from larger Whitewater rivers.
Abundance. COMMON.
Number of specimens examined. 220 from 31 collections. These
specimens are the first to be reported since the unique holotype was
described.
Food. CARNIVORE. This species is mostly piscivorous.
Reproduction. Strategy: probably r2. Spawning is probably annual
and synchronous with the onset of the rains.
Migrations. It is probably capable of long local movements to
avoid drought.
Engraulisoma cf taeniatum Castro 1981
Sardinita Anchoa - Anchovy Tetra
Fig. 117. Map: fig. 118. Couplet 73a.
Generic synonymy
Engraulisoma Castro 1981:135 (type species: Engraulisoma taeniatum
Castro 1981, by original designation).
Specific synonymy
Engraulisoma taeniatum Castro 1981:137 (type locality: Brazil, Mato
Grosso, Poconé, Rio Cuiabá).
Comments. Though Apure specimens agree fairly well with the
original description, this could be a new species since the monotypic
genus was hitherto known only from southern Brazil in the Amazon Basin.
Etymology. ENGRAULI = anchovy, SOMA = body; TAENIATUM = striped.

312
Figure 117. Engrauli soma cf taeniatum.

313
Description
Illustrations. Fig. 117.
Diagnosis. This is the only fish in the Apure drainage that looks
like an anchovy but has the teeth (and adipose fin) of a characid. The
lateral line is decurved and passes within 1-4 scales of the anal-fin base.
The rounded abdomen distinguishes them from small Triportheus.
Size. It grows to about 50 mm SL.
Moroholoqy. The pectoral fins are placed low on the body. The
pelvic fins are entirely anterior to the dorsal fin. The dorsal-fin origin
is anterior to the anal-fin origin. The lateral line is complete.
Counts. DR Ü8; AR Ü22; PR 9; VR i6; LLS 35; TS 8; premaxillary
teeth 2-4; no teeth are visible on the maxilla.
Pigmentation. A wide dark lateral stripe extends from behind the
opercle (which it does not touch, however) to the base of the caudal fin,
but is separate from the spot on the tail. The rest of the body is white
in preservative.
Pistribution and Natural History
Range. Amazon and Orinoco basins.
Apure distribution. Map: fig. 118. The anchovy tetra is widely
distributed from mountains to llanos.
Habitat. This species is found in sections of larger rivers and
streams with fast currents.
Abundance. COMMON.
Number of specimens examined. 129 from 31 collections.
Food. It probably feeds on items of the drift carried down by the
current.
Reproduction. Strategy: probably rl.

Figure 118. Apure drainage distribution of Engraulisoma cf taeniatum.

CO
t—*
cn
CUENCA DEL RIO APURE
UtRlOA ^
V

316
Galeocharax sp.
Dientón Alargado
Fig. 119. Map: fig. 120. Couplet 42b.
Generic synonymy
Galeocharax Fowler 1910:790 (type species: Cynopotamus gulo Cope
1870:565, by original designation); Menézes 1976:40 (diagnosis).
Comments. Menézes (1976) diagnosed the three genera of the
subfamily Cynopotaminae by means of a key. He recognized three spe¬
cies: G. humeralis from the Paraguay and Lower Paraná basins, G. guio
from the Amazon and Sao Francisco basins, and G. kneri from the upper
Paraná. Of these G. humeralis can be eliminated as representing the
Apure specimens since it has 98-101 lateral-line scales. The range of
79-84 LLS seen in Apure specimens agrees with both the remaining spe¬
cies, G. gulo and G. kneri which are separable only on the basis of
morphometries and number of teeth on the dentary. Thus, a careful
comparison with Amazonian material is necessary to place the Apure
specimens, which represent a new record for the Orinoco Basin for the
genus and probably represent a new species as well.
Etymoloqy. GALEO = helmet, from the Greek, CHARAX = palisade,
from the Greek (Eigenmann 1927), in this case a palisade of teeth, but
also given to mean a "gnasher of teeth" in Eigenmann’s earlier works,
Charax is a genus of characid frequently used in the formation of
generic names.
Description
Illustrations. Fig. 119.
Diagnosis. Galeocharax is characterized by the following combi¬
nation of characters: the lower edge of the cleithrum is sinuous, not

317
Figure 119. Galeocharax sp. After Menézes (1976).

318
notched; the anterior dorsal region is only slightly elevated; the
dentary has an inner row of 7-11 small conical teeth; the nasal bone is
laminar; the ectopterygoid is without a ventral bony crest; the anal
fin has about 47 rays; and the lateral line has 80-101 pored scales
(Menézes 1976). This species is easily confused with Acestrocephalus
cf boehlkei, but can be distinguished by the absence of black spots on
the tip of the chin and the anal and dorsal-fin origins (present in
young Acestrocephalus), and in having the first pair of canines widely
separated (by a distance at least as great as their length).
Size. It grows to at least 90 mm SL.
Morpholoov. The scales are ctenoid. The lateral line is com¬
plete.
Counts. DR 11; AR 47; LLS 79-84.
Pigmentation. It is mostly plain silvery with transparent fins.
Pistribution and Natural History
Range. Until now, the genus Galeocharax has been known only from
the Amazon, Paraná and Sao Francisco basins.
Apure distribution. Map: fig. 120. This species is known only
from large rivers in the low llanos.
Habitat. Galeocharax sp. inhabits large Whitewater rivers.
Menézes (1976) hypothesized that Galeocharax may compete with Acestro¬
cephal us in the Amazon basin, and thus restrict the latter to the upper
reaches of the drainages where they occur together. This does not
seem to be the case in the Apure drainage, but perhaps a similar pat¬
tern will emerge once their distribution throughout the Orinoco Basin
is better known.
Abundance. UNCOMMON.

Figure 120. Apure drainage distribution of Galeocharax sp.

CUENCA DEL RIO APURE
OJ
ro
o

321
Number of specimens examined. 306 from 22 collections.
Food. CARNIVORE. Piscivore.
Reproduction. Strategy: probably r2. Spawning is probably annual
and synchronous with the onset of the rains.
Gephvrocharax valenciae Eigenmann 1920
Sardinita - Tetra
Figs. 31 (spur in male) & 121. Map: fig 122. Couplet 11a.
Generic synonymy
Gephyrocharax Eigenmann 1912:23 (type species: Gephyrocharax chocoensis
Eigenmann 1912, by original designation).
Specific synonymy
Gephyrocharax valenciae Eigenmann 1920:11 (type locality: Isla del
Burro, Lake Valencia, Venez.), seen; Pearse 1920:21 (Río Bue, L.
Valencia, Venez.); Eigenmann & Myers 1929:484 (L. Valencia area,
Venez.); Schultz 1944b:323 (key, Barquisimeto, L. Valencia, upper Rio
Guárico, and Caripito, Venez.); Mago L. 1970:70; Géry 1977:353, 358
(key).
Types. Holotype: CAS 44297 (formerly IU 15131); Paratypes: CAS
44294(99) (IU 15129); CAS 44295(2) (IU 15130); CAS 44296(3) (IU 15131).
Etymology. GEPHYR0 = bridge, from the Greek, CHARAX = palisade,
from the Greek (Eigenmann 1927), in this case a palisade of teeth, but
also given to mean a "gnasher of teeth" in Eigenmann’s earlier works,
Charax is a genus of characid frequently used in the formation of
generic names and Eigenmann interpreted Gephyrocharax as a bridge
between the genera Paraqoniates and Flysteronotus; VALENCIAE = after
Lake Valencia, type locality of this species.

322
Figure 121. Gephvrocharax valenciae

323
Description
I11ustrations. Figs. 41 (caudal spur of males) & 121; ?Géry
1977:352 (as unidentified species of Gephyrocharax), 353; Román
1985:158.
Di agnosis. The development of a caudal spur in males (fig. 41) is
known only in this species and Corvnopoma riisei. The latter lacks an
adipose fin (present in G. val ene iae). This species is further
distinguished by the straight dorsal profile, and the posterior position
of the dorsal-fin origin (well behind the anal-fin origin).
Size. This fish grows to about 50 mm SL, but most individuals are
between 30-40 mm.
Morphology. The dorsal-fin origin is midway between a point an eye
diameter behind edge of the opercle and the caudal-fin base. The eye is
longer than the snout, and measures three times in the head length, but
is slightly less than the interorbital width. The frontal fontanel
extends to the ethmoid (middle of the interorbital space).
Counts. DR ii7-ii8; AR 30-32; PR i9-i10; VR i6; LLS 40-42; TS 5-6
1/2 above LL, 5-7 below.
Measurements. Head 4.33, Depth 3.33-3.66.
Pigmentation. The body is silvery, without a humeral spot. There
is a silvery lateral stripe underlain with black pigment that continues
as a diffuse caudal blotch, but which does not extend onto the tips of
the central caudal-fin rays. The jaws are black-tipped.
Pistribution and Natural History
Range. Orinoco Basin.
Apure distribution. Map: fig. 122. It is common throughout the
drainage.

Figure 122.
Apure drainage distribution of Gephyrocharax valenciae.

I08
9°
CUENCA DEL RIO APURE
LO
ro
cn

326
Habitat. It occurs in almost any body of running water, but is
more common in the piedmont.
Abundance. ABUNDANT.
Number of specimens examined. 5,636 from 211 collections.
Food. CARNIVORE. This fish usually feeds on terrestrial in¬
sects, aquatic insects and microcrustaceans (K. Winemiller, pers.
com.). Pearse (1920) reported a similar but more varied diet that
included algae, plants, chironomids, beetles, ants, cladocerans, Cy¬
clops, ostracods, mites, and sediment from Lake Valencia.
Reproduction. Strategy: rl; generation time: 4 months; fecundity:
734 eggs/female; egg diameter: 0.75 mm (Winemiller & Taphorn 1989).
This species spawns during the most of the rainy season. Individuals
may spawn more than once per season.
Importance. Ornamental.
Gnathocharax steindachneri Fowler 1913
Dientoncito
Fig. 123. Map: fig. 124. Couplet 6a.
Generic synonymy
Gnathocharax Fowler 1913b:560 (type species: Gnathocharax steindachneri
Fowler, by original designation).
Specific synonymy
Gnathocharax steindachneri Fowler 1913:561, fig. 19 (type locality:
"Igarapé do Candelaria, Rio Madeira; Rio Madeira 100 east of the
meridian 60 20’; trib. of Rio Madeira near Porto Velho, Brazil") seen;
Myers 1934:5 (near Caicara, Venez.); Bóhlke 1955b: 1; Mago L. 1970:70;
Géry 1977:304, 305, 311 (key).
Comments. Careful analysis of the Apure specimens might reveal
subspecific differences between them and the Amazonian form.

327
Figure 123.
Gnathocharax steindachneri.

328
Etymology. GNATHO = jaws, CHARAX = palisade, from the Greek
(Eigenmann 1927), in this case a palisade of teeth, but also given to
mean a "gnasher of teeth" in Eigenmann’s earlier works, Charax is a
genus of characid frequently used in the formation of generic names;
STEINDACHNERI after Dr. F. Steindachner, Austrian ichthyologist, and
pioneer in South American taxonomic research.
Description
111ustrations. Fig. 123; Bohlke 1955b:14; Géry 1977:304, 305;
Román 1985:180.
Diagnosis. This fish is distinguished by the following combina¬
tion of characters: an incomplete lateral line; the dorsal-fin origin
behind the anal-fin origin; a large mouth with canine teeth; a maxil¬
lary bone that is longer than the vertical diameter of the eye; and an
anal fin with 40 rays or fewer.
Size. It grows to about 80 mm SL, but most individuals are
smaller.
Morphology. The body is greatly compressed, wedge-shaped in
cross section with the point ventral. The ventral edge of the belly is
keeled between the pectoral and ventral-fin bases. There are only 5-6
pored scales in the lateral line. The ventral profile of the head is
swept upward greatly toward the front, giving rise to an extremely
oblique gape. Thus, the lower jaw juts forward beyond the upper.
There are canine teeth present in both jaws. The pectoral fins are
long and falcate and the caudal fin is deeply forked.
Counts. DR ii7; PR i14-i17; AR i ii26-i ii32; VR i7; LLS 35; GR 9-
10.
Measurements. GBD 3.0-3.4 in SL; HL 3.5-3.8 in SL.

Figure 124. Apure drainage distribution of Gnathocharax steindachneri.

CUENCA DEL RIO APURE
(«CALA miUA
100
OJ
co
o

331
Pigmentation. The body is silvery-gray to tan, and the iris of
the eye, opercle and circumorbital bones are also silvery. There is a
large caudal blotch. A thin black line is present on the sides above
the anal-fin base, and along the anal-fin base. The dorsum has a dark
mid-dorsal line.
Distribution and Natural History
Range. It occurs in the Rio Madeira and Rio Negro in Brazil; the
upper Orinoco in Venezuela, and Guyana.
Apure distribution. Map: fig. 124. This species has been found
only in the Aguaro River system in the southeastern corner of the Apure
drainage.
Habitat. It occurs in blackwater streams with aquatic vegetation.
Abundance. RARE.
Number of specimens examined. 2 from 2 collections.
Food. CARNIVORE. It probably feeds on small fishes and insects
that fall to the water’s surface. It seems to be designed for surface
dwelling, as Bohlke (1955b) pointed out.
Reproduction. Probable strategy: rl.
Importance. Ornamental.
Gvmnocorvmbus thayeri Eigenmann 1908
Sardinita Negra - Black Tetra
Fig. 125. Map: fig. 126. Couplet 85b.
Generic synonymy
Gymnocorymbus Eigenmann 1908:93 (type species: Gymnocorymbus thayeri,
by original designation).

332
Figure 125. Gymnocorvmbus thaveri.

333
Specific svnonvmv
Gymnocorymbus thayeri Eigenmann 1908:93 (type locality: Rio Amazonas
from Tabatinga to Gurupa), seen; Fowler 1940:99; 1945:103 (Colombia);
Géry 1977:367, 451, 453-56.
Types. Holotype: MCZ 19242 (indicated as such by Eigenmann
1918:125). Paratypes: MCZ 19272, 20706, 20710, 20720, 20724, 20789,
20804, 20807, 20984, 20973, 20998, 21001, 21007, 21065,
Comments. It is difficult to distinguish between G. thayeri
Eigenmann 1908 and G. socolofi Géry 1972 using Géry’s (1977) criteria.
The Apure drainage population may be the latter species.
Etymology. GYMN0 = naked, from Greek, and means in this case
unsealed, CORYMBUS = summit or nape in this case, also from Greek,
alluding to the unsealed predorsal line in this genus; THAYERI named in
memory of Mr. S. V. R. Thayer, a volunteer of the Nathaniel Thayer
Expedition to Brazil.
Description
II1ustrations. Fig. 125; Géry 1977:453, 456; Axelrod et al.
1971:F-302.00.
Piagnosis. The deep, highly compressed body and the double,
vertically elongated humeral bars are the first clues to the identity
of this species. Although it is superficially quite similar and is
easily confused with Ctenobrycon spilurus, Gymnocorymbus thayeri lacks
the ctenoid scales found in the former. It is distinguished from the
look-alike Poptella orbicularis in lacking the embedded predorsal spine
characteristic of that species, and by the absence of black pigment on
the first anal-fin rays.
Size. It grows to about 50 mm TL.

334
Morphology. The snout is short and the mouth oblique. The lateral
line is complete. The anal fin is relatively long with a nearly straight
margin. The anal and caudal fin are scaled at their bases. The first and
second suborbital are narrow and do not cover the entire cheek (ie. they
don’t extend postero-ventrally to the preopercular bone).
Counts. DR 11; AR 34-41, usually 37-38; LLS 32.
Measurements. HL 3.5 in SL; GBD 1.4-2.1, mean 1.7 in SL.
Pigmentation. The body is silvery, the humeral spot double, and
vertically elongate. All fins are reddish in young fish.
Pistribution and Natural History
Range. It occurs in both the Amazon and Orinoco basins.
Apure distribution. Map: fig. 126. This fish is most commonly
taken from blackwater streams in northern Apure state, from the Aguaro
River system in the southeasternmost part of the drainage, and in
Barinas in the Suripá system.
Habitat. Lowland llaneran streams, usually with clear or black-
water, located in the sandy grassland areas of the drainage are the
typical habitat of this species.
Abundance. COMMON.
Number of specimens examined. 11,035 from 157 collections.
Food. OMNIVORE. It eats a wide variety of small foods, seeds,
insects etc.
Reproduction. Strategy: probably rl. Eggs are scattered among
vegetation and then abandoned. In captivity, live foods are usually
required to bring them into peak condition for spawning. Males have
tiny hooks on the anal fin, that are probably used during fertilization.
Importance. Ornamental.

Figure 126. Apure drainage distribution of Gymnocorymbus thayeri.

336

Hemibrycon metae Myers 1930
Sardina de Montaña - Mountain Tetra
337
Fig. 127. Map: fig. 128. Couplet 91a.
Generic synonymy
Hemibrycon Giinther 1864:318 (type species: Tetragonopterus polyodon
Giinther, bymonotypy).
Specific synonymy
Hemibrycon metae Myers 1930:68 (type locality: Colombia, Guaicaramo, at
the junction of the Rio Guavio and the Rio Upia, almost due east of
Bogotá and north of Barrigón. The elevation is between 300 and 400
meters and there is direct communication with the Meta through the
Upia), seen; Géry 1962:65.
Hemibrycon taeniurus (not Gill) Eigenmann 1920:11 (El Consejo, Río Tuy
and Rio Tiquirito, Venez.).
Hemibrycon dentatus Eigenmann 1927:412 (Villavicencio, Colombia).
Hemibrycon dentatus metae Schultz 1944b:363 (key); Mago L. 1970:71.
Types. Holotype: CAS-SU 23727, examined. I also examined the
holotype, FMNH 56253, of the related H. dentatus.
Comments. Myers (1950) gave the following diagnosis: "Allied to H.
dentatus (Eigenmann) and H. decurrens (Eigenmann) in the sagging lateral
line, differing in the smaller eye, shorter head, and projecting lower
jaw, and variously from one or the other in anal-fin position, depth of
peduncle, and number of scales." He further suggested that given
sufficient material those three species might prove to be only
subspecifically different.
Schultz (1944b) considered all his specimens as subspecies of H.
dentatus. but Géry (1962, 1977) pointed out that H. dentatus is

338
Figure 127. Hemibrycon metae.

339
restricted to the Cauca River and that Schultz’s paratypes included
small specimens of Brvconamericus, accounting for some of the
variability Schultz noted in the original description of his H. dentatus
.jabonero from the Maracaibo Basin.
I have adopted the system supported by Géry (1962, 1967), recog¬
nizing most populations as distinct species.
Etymology. HEMI = half, BRYCON = Brvcon. a genus of characid; METAE
= for the Rio Meta drainage, where the species was first collected.
Description
Illustrations. Fig. 127.
Diagnosis. The jet black stripe through the central caudal-fin rays
is characteristic of this species. It is easily confused with Astvanax
species, but is differentiated by the presence of several (up to 20) teeth
on the maxilla (the number increases with size). Small specimens are simi¬
lar to Brvconamericus species but have more lateral-line scales (42 or
more).
Size. It grows to 100-120 mm SL.
Morphology. The lateral line is complete, a bit decurved anteriorly
and then running low along the sides. The caudal fin is naked except for a
very large scale at the base of each lobe. The dorsal-fin origin is
slightly closer to the tip of the snout than to the of the caudal fin. The
anal fin has a single scale row at its base.
Counts. DR ii8-ii9; AR i i i 26-30; PR i 11-i13; VR Í6-Í8; LLS 42-44; TS
17 8-9 above LL, 6-8 below LL; PDS 13; premaxilla with two rows of teeth, 3-
5 in the outer, four in the inner; maxillary teeth four in smaller speci¬
mens, up to 20 in largest, dentary teeth eight (four large, four smaller);
GR 11.

Figure 128. Apure drainage distribution of Hemibrycon metae.

LA UNIVERSIDAD 0>>E SlCMBE '
UNIVERSIDAD NACIONAL EXPERIMENTAL
DE LOS LLANOS OCCIDENTALES
"EZEQUIE L ZAMORA”
VICE-RECTORADO fcN PORTUGUESA
A .
CUENCA DEL RIO APURE

342
Measurements. GBD 3.0-3.5 in SL. HL 4.66 in SL.
Pigmentation. The dark black lateral stripe is continuous, and runs
from a vague humeral spot posteriorly to the tips of the central caudal
rays. The dorsal adipose and anal fins are speckled with melanophores, and
most of the body is also somewhat flecked with pigment. The tips of the
anal-fin rays are often milk white.
Distribution and Natural History
Range. It occurs in Orinoco Basin.
Apure distribution. Map: fig. 128. This species is most common in
the Andes mountains and piedmont.
Habitat. Andean streams, with fast currents, clear clean water, and
gravel to rock substrate are the typical habitat of this species.
Abundance. COMMON.
Number of specimens examined. 1,190 from 87 collections.
Food. OMNIVORE. This species feeds on insects, seeds, and other
items brought downstream by the current.
Reproduction. Strategy: probably r2. Probably annual and synchronous
with the onset of the rains.
Hemigrammus cf analis Durbin 1909
Sardinita - Tetra
Fig. 129. Map: fig. 130. Couplets 34b & 66a.
Generic synonymy
Hemigrammus Gill 1858:420 (type species: Poecilurichthvs (Hemigrammus)
unilineatus Gill, by monotypy).
Holopristis Eigenmann 1903:145 (type species Tetragonopterus ocellifer
Steindachner 1882:179, by original designation).
Holopristes Eigenmann & Ogle 1907:11 (type species: Tetragonopterus
ocellifer Steindachner 1882:179, by original designation).

343
Figure 129. Hemigrammus cf anal is.

344
Specific synonymy
Hemigrammus analis Durbin 1909:64 (type locality: Rockstone, British
Guiana), seen; Eigenmann 1918:171 (key, synonymy); Géry 1977:371,494
(key).
Types. Holotype: FMNH 53041 (CM1466), examined; Paratypes: FMNH 45
specimens from three localities; CAS-SU 21913(2), 21914(1), 44360(9),
44361(5), 44362(1).
Comments. This identification is tentative.
Etymoloqy. H EM I = half, GRAMMUS = line, referring to the
incomplete lateral line; ANALIS = probably refers to the relatively
short anal fin.
Description
Illustrations. Fig. 129; Eigenmann 1918:pi. 22.
Diagnosis. This species has fewer anal-fin rays (12-15 vs 19 or
more in the other species) than any other Hemigrammus species in the
Apure drainage.
Size. It can grow to about 40 mm SL.
Morphology. As in all Hemigrammus the lateral line is incomplete,
and the base of the caudal fin is scaled. There are about eight pored
scales. They do not extend beyond the tips of the depressed pectoral
fin.
Counts. DR 11; AR 12-15; LS 30-32; TS 5 above, 3 below LL;
premaxilla with 5 seven-cusped teeth in the inner row, 4 in the outer
row (on each side); maxilla with one multicuspid tooth; GR 9.
Measurements. HL 3.5-3.8 in SL; GBD 3.2-3.5.
Pigmentation. The humeral spot is weak and rounded. The lateral
stripe does not extend onto the tail. There is a dark line or dot along
the ventral midline of the caudal peduncle.

Figure 130. Apure drainage distribution of Hemigrammus anal is.

CUENCA DEL RIO APURE
346

347
Distribution and Natural History
Range. It occurs in Guyana, and in the Orinoco basin.
Apure distribution. Map: fig. 130. This species has been found
only in the Aguaro River system drainage, in the southeastern corner of
the Apure drainage.
Habitat. Blackwater streams of low llanos in sandy savanna are
the preferred habitat of this species.
Abundance. UNCOMMON.
Number of specimens examined. 147 from 11 collections.
Food. OMNIVORE. Seeds, microcrustaceans, and insects are the main
items of this species’ diet.
Reproduction. Strategy: probably rl; most Hemigrammus have a
fecundity of 100-200 eggs/female with an egg diameter of about 0.75 mm.
It reaches sexual maturity quickly, in about 4 months, and probably
spawn repeatedly throughout most of the wet season (Winemiller &
Taphorn 1989).
Importance. Ornamental.
Hemigrammus barrigonae Eigenmann & Henn 1914
Sardinita Dos Líneas - Two-Lined Tetra
Fig. 47 (pigmentation) & 131. Map: fig. 132. Couplets 65a & 70a.
Specific synonymy
Hemigrammus barrigonae Eigenmann & Henn 1914:232 (type locality:
Barrigona, Colombia (upper Rio Meta drainage), seen; Eigenmann
1918:161 (key, description); Géry 1977:490, 502, 503, 528.
Types. Holotype: CAS 44368 (originally IU 13423). Paratypes: CAS
44369(62) (IU 13424).

348
Figure 131. Hemiqrammus barrigonae. From Eigenmann (1918).

349
Comments. The variably pored lateral line observed in this species,
which was noted by Eigenmann (1918), has been confirmed in Apure
specimens. See comments under Moenkhausia copei.
Etymology. BARRIGONAE = after Barrigona, Colombia where the type
series was collected.
Description
Illustrations. Fig. 47 (pigmentation) & 131; Eigenmann 1918:pi.
93; Géry 1977:528.
Di agnosis. The deep body and distinctive pigmentation pattern
(figs. 47 & 131) are the best guides to the identification of this
species. The lateral line is usually incomplete, but sometimes can
extend well back behind the dorsal-fin origin. Specimens with a nearly
complete lateral line can be confused with Moenkhausia copei, but are
usually deeper bodied.
Size. It can reach about 40 mm SL.
Morphology. The origin of the dorsal-fin is equidistant from snout
and base of middle caudal rays. The origin of the anal fin is under
that of the dorsal fin. The third suborbital is in contact with the
preopercle behind, leaving no naked area on the cheek.
Counts. DR 11; AR 22-25; LS 33-35; TS 11; maxilla with 4 or 5,
6-pointed teeth.
Measurements. HL 4 in SL; BD 2.6 to 2.8 in SL.
Pigmentation. The base color is light tan to olive. The base of
the anal fin is marked by a black line. A lateral stripe is present
from the operculum to the caudal-fin base, but is not usually continued
onto the tail. Humeral and caudal spots are present but overlaid by
lateral stripe. The outer margin of the anal fin is usually pigmented.

Figure 132. Apure drainage distribution of Hemigrammus barrigonae.

CUENCA DEL RIO APURE
O CAL A iMflCt
OJ
en

352
Distribution and Natural History
Range. It is found in the Orinoco Basin.
Apure distribution. Map: fig. 132. This species occurs in
northern Apure state, in the Aguaro River system of the southeasternmost
part of the drainage, and in Suripá system of Barinas state.
Habitat. H. barriqonae lives in the quieter areas of blackwater
streams with abundant aquatic vegetation, usually in areas with sandy
soils.
Abundance. COMMON.
Number of specimens examined. 1,373 from 57 sites.
Food. OMNIVORE. It eats seeds, microcrustaceans, and insects.
Reproduction. Strategy: probably rl. See account of FL analis.
Importance. Ornamental.
Hemigrammus cf elegans (Steindachner) 1882
Sardinita Elegante - Elegant Hemigrammus
Fig. 133. Map: fig. 134. Couplet 57b.
Specific synonymy
Tetragonopterus elegans Steindachner 1882:179 (type locality: Obidos
(Middle Amazon) Brazil), not seen; Eigenmann & Eigenmann 1891:53.
Hemigrammus elegans Eigenmann 1910:436; 1918:143 (key, description) Géry
1977:510 (key).
Comments. This identification is tentative.
Etymology. ELEGANS = elegant, pretty.
Description
Illustrations. Fig. 133. Eigenmann 1918:pi. 17, fig. 3.
Diagnosis. This is the only Hemigrammus in the Apure drainage with a
black mark in the dorsal fin. However, It might be confused with

353
Figure 133. Hemiqrammus cf elegans.

Figure 134. Apure drainage distribution of Hemigrammus cf elegans.

355

356
Hyphessobrycon bentosi. but that species has a large, well-defined
humeral spot, whereas elegans has none. Megalamphodus cf axelrodi
is the most similar species, but it has conical (vs tricuspid) teeth.
Size. It grows to about 40 mm SL.
Morphology. It is somewhat elongate and compressed.
Counts. DR 11; AR 23-24; LS 30-33; TS 5 above, 3-4.5 below LL.
Pigmentation. The body is pale, with no humeral spot. The dorsal fin
and sometimes the anal fin have a faint blotch of pigment on the first few
rays. The pigment on the anal fin is either quite variable or fades rapidly
in preservative.
Distribution and Natural History
Range. It is known from the Amazon and Orinoco basins.
Apure distribution. Map: fig. 134. It occurs in northern Apure
state and in the Aguaro River system.
Habitat. It inhabits blackwater streams of the low llanos.
Abundance. UNCOMMON.
Number of specimens examined. 50 from 6 collections.
Food. OMNIVORE. It eats seeds, microcrustaceans, and insects.
Reproduction. Strategy: probably rl; see account of H. anal is.
Importance. Ornamental.
Hemigrammus marginatus Durbin-Ellis 1911
Sardinita - Bassam Tetra
Fig. 45 (caudal pattern) & 135. Map: fig. 136. Couplet 60a.
Specific synonymy
Hemigrammus marginatus Durbin-Ellis 1911:159 pi.3, fig. 3 (type
locality: Queimadas, Río Itapicurú [coastal drainage of southern
Brazil, about 39° ll’S]), seen, Eigenmann 1920:10 (Lake Valencia,

357
Figure 135. Hemigrammus marginatus.

358
Venez.) Pearse 1920:12; Eigenmann & Myers 1929:529 (questioned
Eigenmann’s identification of Lake Valencia specimens); Axelrod et al.
1971:F-322.05; Mago L. 1970:71; Géry 1977:442, 502, 521 (key).
Types. Holotype: FMNH 54430 (CM 3053), examined. Ibarra &
Stewart (1987) report that 130 of the paratypes CM 3054-67, 3077, 3082-
84 are also at FMNH.
Comments. Apure drainage specimens are very similar to the holotype,
though the latter has lost all but a trace of the pigment in the tail.
The holotype has the first nine lateral scales pored on left side, but
only seven on the right. Its air bladder is two chambered, with the
first only half as long as the posterior, which extends down almost to
the anus.
Etvmology. MARGINATUS = refers to the black margin of the caudal
fin.
Description
Illustrations. Figs. 40b (caudal pattern) & 135; Durbin Ellis 1911:
pi. 3, #3 (holotype); Axelrod et al. 1971:F-322.05.
Diagnosis. In fresh material the caudal-fin pigmentation (fig. 40b)
is diagnostic for this species when combined with the incomplete lateral
line. It is easily confused (especially in old or faded material) with
Hemigrammus sp. "arriba," but the latter species has dense pigment only
in the upper caudal-fin lobe, and differs further in having fewer
lateral scales (30-31 vs 32-34), fewer transverse scales (9-10 vs 11),
and fewer scales around the caudal peduncle (10 vs 12-13).
Size. This fish grows to about 45 mm SL, but most individuals are
smaller.
Morphology. The lateral line is incomplete, with only about
eight perforated scales. The anal fin has a short sheath of scales

359
anteriorly. The tail has a few scales covering its base, in the larg¬
est specimens reaching more than 1/3 out length of rays.
Counts. DR Ü9-Ü10; AR i i i 19-i i i22; LS 32-34; TS 11; CPS 13;
teeth 4 or 5 in outer and inner row of premaxilla, none on maxilla,
dentary with 4.
Measurements. HL about 27% SL; EYE about 43% HL.
Pigmentation. The dorsal fin is orange, red or yellow. The caudal
fin is often similarly colored near its base but has a wide marginal or
submarginal black band of pigment. The lateral stripe is silvery in
life, leaden in preservative. The scales on the dorsum are outlined in
gray, and the dorsal midline has a concentration of pigment. The belly
is white. Some populations have a curving row of black dots arranged in
groups of four that extends from the opercle out along the first nine
scales of the incomplete lateral line. There is a concentration of
melanophores on the body at the base of the first ten anal-fin rays.
Distribution and Natural Hi story
Range. It occurs from Venezuela to Argentina.
Apure distribution. Map: fig. 136. It is very common throughout
the lower portions of the drainage.
Habitat. It abounds in the quieter sections of streams, and is
also common in standing water.
Abundance. COMMON.
Number of specimens examined. 3,767 from 184 collections.
Food. OMNIVORE. Microcrustaceans, aquatic insects, terrestrial
insects, seeds, and protozoa comprise the major items of this species diet
(K. Winemiller, pers. com.).
Reproduction. Strategy: probably rl. see account of FL anal is.
Importance. Ornamental.

Figure 136. Apure drainage distribution of Hemiqrammus marginatus.


362
Hemigrammus micropterus Meek 1907
Sardinita - Small-finned Tetra
Fig. 51b, c (caudal spot) & 137. Map: fig. 138. Couplet 71b.
Specific synonymy
Hemigrammus micropterus Meek i_n Eigenmann & Ogle 1907:13,15 (type
locality: Los Castillos, Venezuela), seen; Eigenmann 1918:150 (key);
Fowler 1931:408 (Pitch Lake at Guanoco, Venez.); Mago L. 1970:71; Géry
1977:495 (key).
Types. Holotype: CAS 44438 (IU 10802).
Comments. Hitherto known only from the type specimen.
Etymology. MICRO= small, PTERUS = fin.
Description
111ustrations. Figs. 51b, c (caudal spot) & 137. Eigenmann 1918:pis.
18 & 78.
Diagnosis. The large, triangular caudal spot is the distinguishing
feature of this species. The spot is usually not united with the lateral
stripe, but continues to the tips of the central caudal-fin rays.
Size. It grows to about 40 mm SL.
Morphology. The lateral line is incomplete, with the line of 8-10
pored scales reaching back to the tips of the depressed pectoral-fins,
or just reaching insertions of pelvic fins. There are only a few
scales on the caudal-fin base.
Counts. DR 11; AR 19-21; LS 28-33; Premaxilla with 5-6 multicuspid
teeth in inner row, the central 2 asymmetrical, 3-4 in outer row; maxilla
with one 7-pointed tooth.
Measurements. BD 3.2 to 4.0 in SL.
Pigmentation. See diagnosis.

363
Figure 137. Hemiqrammus micropterus.

Figure 138. Apure drainage distribution of Flemigrammus micropterus.

CUENCA DEL RIO APURE
tlCALA IMNCt
co
cr>
en

366
Distribution and Natural History
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 138. This species is most commonly
taken from northern Apure state, from the Aguaro River system in the south-
easternmost part of the drainage, and in Barinas in the Suripá system.
Habitat. Blackwater streams of the sandy-soiled savanna, with
aquatic vegetation are its preferred habitat.
Abundance. COMMON.
Number of specimens examined. 2,096 from 37 collections.
Food. OMNIVORE. It feeds on seeds, microcrustaceans, and in¬
sects.
Reproduction. Strategy: probably rl; see account of H. anal is.
Importance. Ornamental.
Hemigrammus microstomus Durbin 1910
Sardinita - Small-Mouthed Tetra
Fig. 139. Map: fig. 140. Couplet 66b.
Specific synonymy
Hemigrammus microstomus Durbin j_n Eigenmann 1910:436 (type locality:
Rio Amazonas), seen; Mago L. 1970:71; Géry 1977:494, 499.
Etymology. MICRO = small, ST0MUS = mouth.
Types. Holotype: MCZ 20782. Paratypes: MCZ 20782 & 20783 (19),
20840(5), CAS 42680.
Description
Illustrations. Fig. 139; Durbin i_n Eigenmann 1910:pi. 3. Durbin-
Ellis 1911:pi. 8.
Diagnosis. Pigmentation pattern is the best guide to the identity of
this species. The humeral spot is distinctly rounded and somewhat small, a
faint, diffuse lateral stripe extends from it back to the caudal

367
Figure 139. Hemiqrammus microstomus.

368
peduncle. The rest of the fish is plain, silvery to whitish. There
are 2-3 teeth on the maxilla. The lateral line is incomplete, with
about ten pored scales extending posteriorly to just beyond the tip of
the pectoral fin.
Size. It grows to about 40 mm SL.
Morphology, see diagnosis.
Counts. DR 11; AR 21; LS 31-34; TS 5 above, 3 below LL; premaxilla
with 5 seven cuspid teeth in inner row, 2 in outer row; maxilla with 2-3
multicuspid teeth; GR 7.
Measurements. Eye 2.25 in HL, about equal to interorbital; HL 3.6 to
4 in SL; GBD 2.8-3.5 in SL.
Pigmentation. The fins are all hyaline.
Pistribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map: fig. 140. It has so far been found only in
the Aguaro River system in the southeastern corner of the Apure River drain¬
age.
Habitat. Blackwater streams of the savanna, with aquatic vegetation
are this species primary habitat.
Abundance. UNCOMMON.
Number of specimens examined. 594 from 16 collections.
Food. OMNIVORE. It feeds on a variety of foods including seeds,
microcrustaceans, and insects.
Reproduction. Strategy: Although no data are available this
species is probably an rl strategist like most small tetras. Most
Hemigrammus spp. are egg scatterers, see the account for H. anal is.
Importance. Ornamental.

Figure 140. Apure drainage distribution of Hemigrammus microstomus.

CUENCA DEL RIO APURE —~
CO
O

371
Hemigrammus cf mimus Bohlke 1955
Sardinita Mima - Mimic Tetra
Fig. 46 (caudal pattern) & 141. Map: fig. 142. Couplet 63a.
Specific synonymy
Hemigrammus mimus Bohlke 1955a:229 (type locality: rapids at Camanáos on
the Rio Negro, Brazil (down river from Sao Gabriel); Géry 1977:490.
Types. Holotype: CAS 47759. Paratypes CAS 47760(1).
Comments. I have called this species H. mimus because Apure
drainage specimens share the color pattern of that species. However,
the teeth and anal-fin ray counts differ. Subsequent study may show
them to be related to H. ervthrozonus.
Etymology. MIMUS = mimic, referring to the similarity between this
species and the species of Microschemobrvcon, with which Bohlke
originally confused it.
Description
Illustrations. Fig. 46 (caudal pattern) & 141; Bohlke 1955:232.
Diagnosis. The small embedded crescent of pigment at the base of
each caudal lobe and the small size are both diagnostic of this species.
Microschemobrycon casiquiare share this pigment pattern, but have the
premaxillary teeth in a single row, and is more slender (GBD 3.7-4.6 in
SL).
Size. This is a tiny species which seldom exceeds 30 mm SL.
Morphology. The body is nearly subcylindrical with the greatest
body depth found at the dorsal-fin origin. Males have minute hooks on
the anal and caudal-fin rays.
Counts. DR ii9; AR 17 in type series, but 19 in Apure specimens;
PR ill; VR i7; LS 32, lateral line pores 9-11; TS 8; predorsal scales

372
Figure 141. Hemigrammus cf mimus.

Figure 142. Apure drainage distribution of Hemigrammus cf mimus.

374

375
8-9; premaxillary teeth in two rows, inner with 5-6, outer with 2-3
teeth (said to have but a single tooth in type series in this outer
row), 3-4 slender tricuspid teeth on maxilla; GR 10-11.
Measurements. GBD 3.5-3.6 in SL; HL 3.7 in SL; EYE 2.8-2.9 in HL.
Pigmentation. The body is tan, with the embedded pigment at the
base of each caudal lobe (fig. 41) the only marking.
Distribution and Natural History
Range. This species occurs in the Rio Negro in Brazil, the upper
Rio Orinoco Basin in Venezuela and Colombia.
Apure distribution. Map: fig. 142. This species is found only in
northern Apure state and Barinas.
Habitat. H. cf mimus occurs in sandy savannas, in clear and
blackwater streams.
Abundance. RARE.
Number of specimens examined. 125 from 4 collections.
Food. OMNIVORE. It eats seeds, microcrustaceans, and insects.
Reproduction. Strategy: probably rl; see account of H. anal is.
Hemigrammus newboldi (Fernández Y.) 1949
Sardinita Manchada - Spotted Tetra
Fig. 49 (caudal spot) & 143. Map: fig. 144. Couplet 68a.
Specific svnonvmv
Ramirezella newboldi Fernández Yépez 1949:1-3 (type locality: Pal i tal,
Guárico state, Venezuela), seen; Mago L. 1970:71; Géry 1977:458, 487.
Comments. The description of a new genus for this species seems
totally unwarranted. There are no characters that justify the creation of
Ramirezella, and so it should be considered as a synonym of Hemigrammus.
However, placement of H. newboldi could be with either Hemigrammus or

376
Figure 143. Hemiqrammus newboldi.

377
Moenkhausia. As Eigenmann (1918) pointed out, specimens of Moenkhausia
continho show a complete gradation from an incomplete to a complete lateral
line. Both Hemiqrammus and Moenkhausia, have scaled caudal fins. Fernández
Y. (1949) stated that the basal fourth of the caudal fin is scaled in FL_
newboldi. Géry (1977) pointed out that this species is very similar to
Moenkhausia cotinho, and I agree. It is quite possible, then, that H.
newboldi is not distinct at the specific level either, but rather is a
junior synonym of M. cotinho. A direct comparison with specimens of
that species from Guyana is necessary to make the final determination.
Etymology. NEWBOLDI = after Mr. Philip Newbold, a friend of A.
Fernández Y., who died while working on experiments on Lake Maracaibo,
in 1948.
Description
Illustrations. Fig. 49 (caudal spot) & 143; Fernández Y. 1949:
fig 1.
Diagnosis. The large caudal spot (covering most of the caudal pedun¬
cle and at least the basal third of the caudal fin) is characteristic of
this species.
Size. It grows to about 50 mm SL.
Morphology. The lateral line is incomplete, with 8-10 pored scales
that extend posteriorly to just beyond the pelvic-fin insertions.
Counts. DR ii9; AR iiil8-iiil9; LS 28-30; TS 10; 5 teeth in inner
row of premaxilla, 4 in outer;
Measurements. HL 4.0 in SL; GBD 2.8 in SL.
Pigmentation. In preservative, it is tan with a black lateral stripe
starting under the dorsal fin. The caudal peduncle and base of tail have a
very large black blotch that usually covers most of the caudal peduncle and
at least the basal third of the caudal fin. The humeral spot is diffuse.

Figure 144. Apure drainage distribution of Hemigrammus newboldi.

CUENCA DEL RIO APURE

380
Distribution and Natural History
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 144. So far this species has been
found only in the Aguaro River system in the southeastern corner of the
Apure drainage.
Habitat. Blackwater streams with aquatic vegetation are its
primary habitat.
Abundance. UNCOMMON.
Number of specimens examined. 252 from 23 sites.
Food. OMNIVORE. It eats seeds, microcrustaceans, and insects.
Reproduction. Strategy: probably rl; see account of H. anal is.
Importance. Ornamental.
Hemiqrammus rhodostomus Ahl 1924
Borachito - Rummy-Nosed Tetra
Fig. 45a (caudal pattern) & 145. Map: fig. 146. Couplet 59a.
Specific synonymy
Hemiqrammus rhodostomus Ahl 1924 (type locality: Pará, Brazil) seen;
Ramirez (1960); Géry 1977:502, 520, 547, 550; Axelrod et al. 1971:F-
324.00; Mago L. 1970:71.
Comments. The Venezuelan specimens could represent a new species,
since they differ slightly in color (Apure drainage and Capanaparo
drainage specimens have an orange to red caudal peduncle and caudal fin
in life) and have one small maxillary tooth. To be certain, however,
they will have to be compared with material of H. rhodostomus from other
basins, as well as H. bleheri Géry & Mahnert 1986 from the middle Rio
Negro in Brazil.
Etymology. RHODO = red; ST0MUS = mouth.

381
Figure. 145. Hemigrammus rhodostomus.

382
Description
Illustrations. Fig. 45 (caudal pattern) & 145; Axelrod et al.
1971:F-324.00 Géry 1972:31, 90, 92, 93, 94; 1977:520; Géry & Mahnert
1986:44.
Diagnosis. The distinctive pigmentation of this species
immediately identifies them. The nose is red, and the caudal fin is
marked with white and black bands (fig. 45A).
Size. It grows to about 50 mm TL.
Morphology. The body is slender, and not too compressed.
Counts. DR ii9; AR i i i 12 -13 ; PR ill-12; VR i7; LS 32-33 (9
pored); TS 9.
Measurements. GBD 26-28% SL.
Pigmentation. The body is light tan in preservative, silvery to
greenish in life. The upper and lower edges of the caudal peduncle are
marked with a black line or spot. The snout is blood-red. The caudal
fin is most distinctive, with white lobes crossbanded with black (fig.
45a).
Distribution and Natural History
Range. It has been found in the lower Amazon region, around
Aripiranga and Para, and in the Orinoco Basin.
Apure distribution. Map: fig. 146. It is found only in the Aguaro
River system in the southeastern corner of the Apure drainage.
Habitat. It inhabits blackwater savanna streams, in areas with
sandy soils.
Abundance. RARE.
Number of specimens examined. 116 specimens, 4 collections.
Food. OMNIVORE. It eats seeds, microcrustaceans, and insects.

Figure 146. Apure drainage distribution of Hemigrammus cf rhodostomus.

10*
9"
LA uNlvenSlOAP O'It SICMBF
UNIVERSIDAD NACIONAL EXPERIMENTAL
DE LOS LLANOS OCCIDENTALES
"EZEOUIEL ZAMORA'
VICE RECTOR ADO EN PORTUGUESA
CUENCA DEL RIO APURE
IIMit iiurict
384

385
Reproduction. Strategy: probably rl. Pal icka (1989) reported on thi s
species’ unusual spawning behavior. H. rhodostomus do not scatter eggs in
the usual characid fashion, but rather hang their adhesive eggs from plants
near the surface. Water chemistry was of utmost importance to induce spawn¬
ing also for the successful development of the embryos, and fry. They
require low ph (5-6) soft (about 40 mS conductivity) water stained to tea-
color with peat moss, and from which most calcium has been chemically
removed. No sort of water contamination is tolerated by this species.
They will go into shock and die if water conditions are allowed to deterio¬
rate. Light must be reduced to a minimum. Spawning occurs at night, eggs
are transparent and about 1 mm in diameter. A pair can deposit from 400-
500 eggs per spawning. The fry are rather large, about 3.9 mm when they
have absorbed the yolk, and can immediately feed on brine shrimp nauplii.
They grow quickly, reaching 10 mm in one week if properly fed. They mature
in six to seven months.
Importance. It is highly valued as an ornamental.
Hemigrammus cf schmardae (Steindachner) 1882
Sardinita - Tetra
Fig. 51a (caudal spot) & 147. Map: fig. 148. Couplet 71a.
Specific synonymy
Tetragonopterus schmardae Steindachner 1882:37 pi. vii, fig. 6 (type
Locality: Tabatinga, Brazil), not seen.
Hemigrammus schmardae Fowler 1906:335 (Peruvian Amazon) not seen;
Eigenmann 1910:436; 1918:163 (key); Géry 1977:506, 524, 601.
Comments. Eigenmann (1918) did not mention a black line along the
base of the anal fin. This character is so obvious that it would hardly
have been overlooked if present in his Amazonian specimens. His figure

Figure 147. Hemigrammus cf schmardae.

387
also represents a much deeper-bodied fish than those seen in the Apure
drainage. I suspect, therefore, that my identification of this species
is incorrect. It probably represents a new species.
Etymology. SCHMARDAE = origin unknown.
Description
Illustrations. Fig. 51a (caudal spot) & 147. Eigenmann 1918:pis.
20, 78; Géry 1977:524.
Diagnosis. The small size, relatively large rounded caudal spot,
the absence of a humeral spot, and the black line along the base of the
anal fin are the best clues to the identity of this species.
Size. It is a small species that only rarely reaches 40 mm SL.
Morphology. The lateral line is incomplete, with about 12 pored
scales that extend posteriorly to over the middle of the depressed
pelvic fins.
Counts. AR 20; LS 31; premaxilla with 5 pentacuspid teeth in
inner row, 3 in outer; maxilla with 2 pentacuspid teeth.
Measurements. BD 3.4 to 4.0 in SL.
Pigmentation. There is no humeral spot. The base of the anal
fin is marked with a black line. The caudal spot is quite large,
covering most of peduncle (fig. 51a).
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map: fig. 148. This species has been found
only in the Aguaro River system in the southeastern corner of the Apure
drainage.
Habitat. H. cf schmardae inhabits blackwater savanna streams with
aquatic vegetation.
Abundance. UNCOMMON.

Figure 148. Apure drainage distribution of Hemigrairmus cf schmardae.

389

390
Number of specimens examined. 607 from 16 collections.
Food. OMNIVORE. It eats small seeds, microcrustaceans and
insects.
Reproduction. Strategy: probably rl; see account of H. analis.
Importance. Ornamental.
Hemiqrammus so. "arriba"
Sardinita - Tetra
Fig. 45c (caudal pattern) & 149. Map: fig. 150. Couplet 60b.
Comments. This species is very similar to, but clearly distinct from
H. marqinatus. It is also similar to H. rodwavi, a common species in
eastern Venezuela, but that species has the lateral stripe continuing onto
the central caudal rays without interruption, and also has very different
nuptial coloration. The Apure specimens probably are a new species.
Description
Illustrations. Figs. 45c (caudal pattern) & 150.
Diagnosis. This fish looks like a Hemiqrammus marqinatus which has
black pigment only in the upper lobe of the caudal fin. The eye is rela¬
tively smaller than in H. marqinatus. See the diagnosis for that spe¬
cies.
Size. It reaches about 40 mm SL.
Morphology. The lateral line is incomplete, and has about eight
pored scales. The anal fin has a short sheath of scales anteriorly.
Counts. DR ii9; AR ÍÜ20-22 ; PR 10-11; VR i6; LS 30-31; TS 9-10;
CPS 10; teeth variable on premaxilla, 3 to 5 in outer row, 2 to 5 in inner,
none on maxilla.
Pigmentation. Some specimens have yellow near base of the caudal fin.
The upper lobe of the caudal fin is marked with an upswept black bar (fig.
149), (the caudal fin is actually finely peppered with tiny melanophores

391
Figure 149. Hemigrammus sp. "arriba".

Figure 150. Apure drainage distribution of Hemigranimus sp. "arriba".

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393

394
that are most dense in the dorsal lobe). The dark lateral stripe does
not continue onto the caudal fin. There is an irregular patch of small
black spots on the body over the origin of the anal fin. Black and yellow
pigment is concentrated along the rays of the anal fin.
Pistribution and Natural History
Range. It is apparently limited to the Orinoco Basin.
Apure distribution. Map: fig. 150. It is common in the llanos.
Habitat. It inhabits the lowland llanos and piedmont streams and
ponds.
Abundance. COMMON.
Number of specimens examined. 4,733 from 149 collections.
Food. OMNIVORE. It eats aquatic and terrestrial insects, seeds,
and microcrustaceans.
Reproduction. Strategy: probably rl; see account of H. anal is.
Importance. Ornamental.
Hemigrammus stictus (Durbin) 1909
Sardinita Flamante - Flaming Tetra
Fig. 151. Map: fig. 152. Couplet 65b.
Specific synonymy
Hyphessobrvcon stictus Durbin 1909:71 (type locality: Lama Stop-Off,
British Guiana), seen; Eigenmann 1910:437; 1912:342; 1921:216; Bohlke
1953a:25.
Hemigrammus stictus Géry 1977:490, 491, 505, 508 (key).
Types. Holotype: FMNH 52798 (CM 1197). Paratypes: CAS-SU 21923(3)
21922(2); 146 from 5 collections in FMNH.
Comments. I have not compared Apure material with specimens from
Guyana.

395
Figure,
tion of
life.
151. Hemigrammus
the body amd the caudal
stictus. The stippling in the posterior por-
fin indicates the area that is red in

396
Etymology. STICTUS = marked or spotted, probably in reference to the
distinct humeral spot.
Description
Illustrations. Fig. 151; Eigenmann 1918:pi. 22 fig. 1; Géry 1977:505,
508.
Diagnosis. The brill iant scarlet to cherry-red color on the posterior
portion of the body is diagnostic for freshly preserved or live specimens.
The humeral spot is round, intense, and often surrounded by a pale area.
There is no black caudal spot. It has more pored scales in the lateral line
(about 15) than most species of Hemigrammus.
Size. It can reach about 40 mm SL.
Morphology. The lateral line is incomplete, with about 15 pored
scales that extend almost to the pelvic-fin origins.
Counts. DR 11; AR 26-31; LS 33-35; TS 6 above LL, 4 below; PDS 9-11;
premaxilla with 5 teeth in inner and 2-3 in outer row; maxilla with 2 wide
multicuspid teeth; GR 11.
Measurements. HL 3.5-3.8 in SL; GBD 2.75-3.25; EYE 2.25 in HL; snout
one half of eye; interorbitals less than the eye, about 2.5 in HL.
Pigmentation. See diagnosis. The body is silvery to whitish. The
lateral stripe is very thin and does not reach the caudal-fin. The lobes of
the caudal, dorsal and anal fins are yellowish.
Pistribution and Natural History
Range. It is known from Guyanan rivers, the Orinoco Basin and the Rio
Guainia (upper Rio Negro) in Venezuela.
Apure distribution. Map: fig. 152. It has been found only in the
Aguaro River system in the southeastern corner of the Apure drainage.

Figure 152. Apure drainage distribution of Hemigrammus Stictus.

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399
Habitat. H. stictus inhabits blackwater savanna streams with aquatic
vegetation.
Abundance. RARE.
Number of specimens examined. 1 specimen.
Food. OMNIVORE. It eats seeds, microcrustaceans, and insects.
Reproduction. Strategy: probably rl; see account of H. anal is.
Importance. Ornamental.
Heterocharax macrolepis Eigenmann 1912
Sardinita dientudo
Fig. 153. Map: 154. Couplet 9a.
Generic synonymy
Heterocharax Eigenmann 1912:405. (type species: Heterocharax macrolepis,
by original designation).
Specific synonymy
Heterocharax macrolepis Eigenmann 1912:406 (type locality: Rockstone,
British Guiana), seen; Géry 1977:309, 314.
Types. Holotype: FMNH 53669 (formerly CM 2142), examined. Paratypes:
FMNH 53670 (CM 2143a), FMNH 53671 (CM 2144a), CAS 61482(1) all examined.
Etymology. HETERO = different, CHARAX = a genus of characid; MACRO =
large, LEPIS = scales.
Description
Illustrations. Fig. 153; Géry 1977:309, 314.
Range. It is found in Guyana, in the Rio Negro drainage and the Middle
Amazon basin, and in the Orinoco Basin.
Diagnosis. This fish is distinguished by the following combination of
characters: the anal-fin origin is under the dorsal-fin origin; the maxilla
is well toothed, with conical teeth; the premaxilla has a single row of

400
Figure 163. Heterocharax macro!epis.

401
conical teeth, some of them canines, the dentary has two rows of coni¬
cal teeth (the second consists of very small teeth). The color pattern
is also distinctive (fig. 153). Géry (1977) further characterized them
as follows: preopercular angle without a real spine; opercle not spi¬
nous; clavicle notched; predorsal line scaled; lateral line complete;
first pectoral ray soft; a single series of teeth on upper jaw.
Size. It can reach 50 mm SL, but most individuals are smaller.
Morphology. In Apure drainage specimens the maxilla always reaches
the end of the first suborbital bone. This character may vary, however, as
in at least one paratype the maxilla did not reach the first suborbital.
The snout tip to end of maxilla distance reaches to behind the center of
the eye. The scales on the dorsum are much smaller than those below the
lateral line. In one paratype, there are lateral extensions of the main
lateral line canal. The dorsal-fin origin is almost equidistant between
the snout tip and the caudal-fin base. The pectoral fins reach the pel-
vies, but the pelvics do not quite reach the anal fin. The preopercle is
without a really sharp spine. The eye is quite large, much longer than
snout. The jaws all possess some canine teeth.
Counts. DR ii9; AR 37;PR i12; VR i7; LS 35; TS 13; GR 6 + 4-5 rudi¬
ments on the lower arch. The premaxilla has about 10 conical teeth plus 3-
4 enlarged canines; the maxilla is completely toothed with over 30 conical
teeth, the dentary has 2-3 large canines and numerous smaller conical
teeth.
Measurements. PDL about 49% SL; HL about 27% SL; SNT about 22% HL,
much shorter than eye diameter which is about 48% HL; GBD about 36% SL.
Pigmentation. A wide, distinct lateral stripe is present from opercle
to tail. The dorsal midline is marked with a dark line, as is the anterior
anal-fin base. Another dark line begins on the sides above the anal-fin

I*
Figure 154. Apure drainage distribution of Heterocharax macrolepis

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404
origin and runs back to the caudal peduncle. All fins are clear except the
caudal, which is dusky gray.
Distribution and Natural History
Apure distribution. Map: fig. 154. It has been found only in the
Aguaro River system in the southeastern corner of the Apure drainage.
Habitat. It is restricted to blackwater streams and morichales.
Abundance. RARE.
Number of specimens examined. 5 from 2 collections.
Food. CARNIVORE. It probably feeds on aquatic insects, microcrusta¬
ceans and small fishes.
Reproduction. Probable strategy: rl. It may spawn throughout the
rainy season.
Importance. It is potentially valuable as an aquarium fish.
Hyphessobrvcon bentosi Durbin 1908
Sardinita - Flag Tetra
Fig. 155. Map: fig. 156. Couplet 56a.
Generic synonymy
Hyphessobrvcon Durbin 1908:100 (type species: Hemiqrammus compressus
Meek 1904, by original designation).
Specific synonymy
Hyphessobrvcon bentosi Durbin j_n Eigenmann 1908:101 (type locality:
Obidos, Brazil [Middle Amazon basin]), seen; Eigenmann 1910:436;
1918:183 (key); Géry 1977:479, 483, 500 (key).
Types. Holotype: CAS 42682. Paratypes: MCZ 20842(21).
Comments. The taxonomic distinction between Mega!amphodus sweglesi
Géry 1961 and Hyphessobrvcon bentosi Durbin 1908 is "nebulous," as indicated
by Géry (1977). The premaxillary row of teeth could be interpreted as

405
Figure 155. Hvphessobrycon bentosi.

406
"single" if you accept a few (1-2, usually) as out of line (perhaps in the
process of being replaced), but in some fish it seems that there are two
definite rows. Since this problem cannot be resolved here, I have opted to
use Hyphessobrvcon bentosi which is the older name and thus the one having
priority should these two forms prove to be identical.
Etymology. HYPHESSO = smal1er, from the Greek, BRYCON = "gnasher of
teeth" in Greek, and also a genus of characid; BENTOSI = after Sr. Bentos,
the collector of the type series.
Description
Illustrations. Fig. 155 (a large adult is depicted in the figure,
most specimens are not so robust); Eigenmann 1918:pis. 25, 79; Géry
1977:483.
Diagnosis. The large black mark in the dorsal fin, and large distinct
humeral spot are characteristic of this species. The premaxilla has two
rows of teeth. The anal fin has 26-27 rays.
Size. It grows to about 30 mm SL.
Morphology. The lateral line is incomplete.
Counts. DR 11; AR iii23-iii27; LS 25; TS 5-6/3-4; 7-9 teeth in inner
row of premaxilla; GR 14.
Measurements. HL 3.3-3.5 in SL; GBD 2.8-3.2 in SL.
Pigmentation. The holotype has lost the humeral spot if one was
present. In Apure drainage specimens, the dorsal fin has a distinct black
spot and a dark humeral spot is present, but there is no caudal spot.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map: fig. 156. It is known only from the low
11 anos.

Figure 156. Apure drainage distribution of Hyphessobrycon bentosi.

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409
Habitat. H. bentosi occurs in quiet blackwater llaneran streams and
is usually associated with fine-leaved aquatic vegetation.
Abundance. UNCOMMON.
Number of specimens examined. 112 from 21 collections.
Food. OMNIVORE. It eats insects, microcrustaceans, and seeds.
Reproduction. Strategy: rl. It spawns throughout the wet season.
Importance. Ornamental.
Hyphessobrvcon metae Eigenmann & Henn 1914
Sardinita del Meta - Meta Tetra
Figs. 50 & 157. Map: fig. 158. Couplet 69a.
Specific synonymy
Hyphessobrvcon metae Eigenmann & Henn 1914:233 (type locality: Rio Meta
at Barrigona, Colombia), seen; Eigenmann 1918:203 (key); Géry
1977:469, 471, 472 (key).
Types. Holotype: CAS 61751 (IU 13421); Paratypes: CAS 61752(2) (IU
13422).
Etymology. METAE = from the Rio Meta.
Description
Illustrations. Figs. 50 & 158; Eigenmann 1918:pi. 93; Géry 1977:
469.
Diagnosis. The wide dusky lateral band and overall darkly pigmented
body are characteristic of this species. There are no marks in the dorsal
or anal fins. The lateral line is short, with only 6-7 pored scales.
Size. It reaches about 40 mm SL.
Morphology. The lateral line is incomplete with about six pored
scales extending to middle of the depressed pectoral fins. The maxilla
does not reach back to the level of pupil. There is a row of scales
along the base of the anal fin; the caudal-fin base is also well scaled.

410
Figure 157. Hvnhessobrvcon metae. From Eigenmann (1918).

Figure 158. Apure drainage distribution of Hyphessobrycon metae.

0««» i o O IOO
CUENCA DEL RIO APURE 1 "
412

413
Counts. DR ilO; AR i i i 18-i i i 20, hooks on first 6 rays; PR i 9; VR i 7;
LS 28-32 PDS 10 TS 9; premaxilla with 2 teeth in outer row, 6 in inner, 1-2
teeth on maxilla.
Measurements. HL 3.5-4 in SL.
Pigmentation. The head, body and fins are all heavily pigmented with
melanophores. The pigment in the caudal fin is concentrated along the
central rays. A pale unpigmented area is present just above the posterior
margin of the anal-fin base. The dorsal, adipose, upper and lower caudal-
fin lobes have red patches in some populations (perhaps varying with
season). The holotype has a triangular spot on the central caudal-fin
rays.
Distribution and Natural History
Range. It is known from the Orinoco Basin.
Apure distribution. Map: fig. 158. It is most common in northern
Apure state and the Aguaro River system.
Habitat. It is found in blackwater streams and morichales.
Abundance. COMMON.
Number of specimens examined. 348 from 28 collections.
Food. OMNIVORE. Microcrustaceans, seeds, and insects comprise its
diet.
Reproduction. Strategy: rl. It probably spawns throughout the rainy
season.
Importance. Ornamental.
Iguanodectes soilurus (Giinther) 1864
Sardinita - Spindle Tetra
Fig. 42 (maxilla) & 159. Map: fig. 160. Couplet 48a.
Generic synonymy
Iguanodectes Cope 1872:260 (type species: Iguanodectes tenuis Cope).

414
Figure 159. Iauanodectes spilurus.

415
Specific synonymy
Piabucus spilurus Giinther 1864:344 (type locality: Cupai River);
Eigenmann & Eigenmann 1891:57; Steindachner 1891:23; Eigenmann
1910:440.
Iguanodectes spilurus Mago L. 1970:71; Géry 1970c:417; 1977:368, 368,
374.
Etymology. IGUANO = from "Iguana, native Haytian name of a lizard"
(Cope 1871), DECTES = a receiver (allusion obscure); SPIL = spot, URUS =
tail.
Description
Illustrations. Fig. 42 (maxilla length) & 159; Géry 1970:figs. 2, 5;
1977:368, 369.
Diagnosis. The teeth of this species are slightly asymmetrical,
greatly expanded (widened) at the tip, and multicuspid. The premaxilla has
an inner row of five teeth, but the outer has only one or two. The body is
long and slender, with a high lateral-line scale count (54-64).
Size. It can grow to about 90 mm SL, but most individuals are between
40-60 mm.
Morphology. This is an elongate, spindle shaped fish with a very
small mouth that is entirely in front of the eye. The dorsal fin is poste¬
rior in position, with its origin behind the midpoint of the SL, but
anterior to the origin of the anal fin. The gill membranes are united,
but free from isthmus. The breast and preventral area are rounded.
The lateral line is complete. The eye is longer than the snout. Vari
(1977) described the unique anatomy of the three chambered swim bladder
in this species and presented characters in support of the monophyly of
the subfamily Iguanodectinae.

Figure 160, Apure drainage distribution of Iguanodectes spilurus.

I»C«U lairict
I—*

418
Counts. DR ii8-ii9; AR iii31-iii36; PR i 12-i13; VR i7; LLS 54-64; TS
6-8 above LL, 4-5 below. The premaxilla has 5-6 multicuspid teeth that are
constricted at their bases and expanded at the tips, maxilla with 1-2 teeth,
dentary with 7-8.
Measurements. HL 4.3-4.7 in SL; GBD 4.5-5.0 in SL.
Pigmentation. The tail is marked with a large black blotch. There is
a thin black line along the posterior lateral midline and another above the
base of the anal fin.
Distribution and Natural Hi story
Range. It occurs in the Amazon Basin in Brazil and Peru, in Guyana,
and in the Orinoco Basin.
Apure distribution. Map: fig. 160. This species is known from north¬
ern Apure state and the Aguaro River system.
Habitat. It frequents blackwater streams, and is usually seen in the
current.
Abundance. UNCOMMON.
Number of specimens examined. 307 from 20 collections.
Food. OMNIVORE.
Reproduction. Strategy: rl. It probably spawns throughout the rainy
season. Males develop hooks and lappets (a hook-like structure with the
free end directed proximally, toward the body of the fish) on the anterior
anal rays during the reproductive season.
Importance. Ornamental.
Markiana geavi (Pellegrin) 1908
Percha - Red-Bellied Tetra
Fig. 161. Map: fig. 162. Couplet 39a.

419
Figure 161. Markiana qeayi.

420
Generic synonymy
Markiana Eigenmann 1903:145 (type species: Tetraqonopterus nigripinnis
Perugia 1891:643, by original designation).
Specific synonymy
Tetraqonopterus (Markiana) geayi Pellegrin 1908:347 (type locality:
Apure, Venezuela), seen.
Markiana geayi Eigenmann 1910:438; 1918:124.
Markiana nigripinnis Mago L. 1970:71; Román 1985:178 (Venez.).
Comments. Although Pellegrin (1908) commented that M. geayi lacked
the caudal spot present in M. nigripinnis from the La Plata Basin, his
description was based on only one specimen. Some Apure specimens lack
caudal pigment, but most have a large black spot on the caudal base; I
suspect that Pellegrin’s specimen had simply lost its color. This then
raises the question of synonymy since the two agree fairly closely in the
few counts and measurements given by Pellegrin. Since I cannot make a
direct comparison, I have chosen to maintain the name M. geayi since it is
based on specimens from the Apure drainage. In most cases, similar species
from northern and southern South America have ultimately proven to be dif¬
ferent, a factor related largely to the large geographic distances involved.
Even so, unless these two forms differ in characters other than those used
so far to distinguish them, the southern and northern forms would appear to
be synonyms.
Etymology. MARKIANA = was named after a teacher of C. Eigenmann, Dr.
Edward Laurens Mark, who for more than 25 years was head of the Harvard
University Zoology Department, where Eigenmann was a student; GEAYI = is
named after F. Geay, a contemporary of Pellegrin who reported on the fisher¬
ies of the Orinoco Basin in his work "Peches dans les Affluentes de l’Ori-
noque" (Schultz 1944b).

421
Description
Illustrations. Fig. 161; Román 1985:178.
Diagnosis. The crenate scales that gradually become smaller in the
rows below the lateral line, the dark horizontal wavy lines on the sides,
the red breast (in life), and scaled anal and caudal fins are characteristic
of this species.
Size. It grows to about 90 mm SL.
Morphology. The lateral line is complete, the anal fin and caudal fin
are scaled, the gill rakers are setiform, and the dorsal-fin origin is
posterior to the pelvic-fin origin.
Counts. DR Ü9-Ü10; AR iii37-iii40; PR i 11-i 12; VR i7; LLS 41; TS
8-9 above LL, 7-8 below.
Pigmentation. The body is silvery with wavy lines of melanophores
between the scale rows on the sides. The pectoral, pelvic and anal fins as
well as the breast are red. The humeral spot is diffuse. A black caudal
spot is present, but not obvious in all preserved specimens.
Distribution and Natural Hi story
Range. It occurs in Orinoco Basin.
Apure distribution. Map: fig. 162. It occurs throughout the
llanos.
Habitat. It inhabits lentic and lotic systems in the llanos.
Abundance. COMMON.
Number of specimens examined. 878 from 124 collections.
Food. OMNIVORE. Its diet includes seeds, terrestrial insects, and
aquatic vegetation.
Reproduction. Strategy: r2; fecundity: 3398 eggs/ovary; egg diame¬
ter: 1 mm. This species spawns once a year, during the first months of the
rainy season. There is no parental care. (Winemiller & Taphorn 1989).

Figure 162. Apure drainage distribution of Markiana neayi

423

424
Importance. Ornamental.
Meqalamphodus cf axelrodi (Travassos) 1959
Sardinita - Calypso Tetra, Red Pristella
Fig. 163. Map: fig. 164. Couplet 57a.
Generic synonymy
Meqalamphodus Eigenmann 1915b:49 (type species: Meqalamphodus
meqalopterus Eigenmann 1915b:150, by original designation).
Ectrepopterus Fowler 1943b:313 (type species: Meqalamphodus
uruouavensis Fowler 1943b:313, by original designation).
Specific synonymy
Aphyocharax axel rodi Travassos i_n Axelrod 1959:3 (type locality:
Trinidad), not seen.
Meqalamphodus axelrodi Axelrod et al. 1971:F-48.00; Géry 1977:572, 586.
Etymology. MEGALAMPHODUS is from Greek meaning "with spacious ways"
(Eigenmann 1919); AXELRODI = after Dr. Herbert Axelrod.
Description
Illustrations. Fig. 163; Géry 1977:572; Axelrod et al. 1971:F-
48.00.
Diagnosis. In life the body is mostly red posteriorly and the dorsal
fin has a black blotch on the upper margin of the anterior rays. The teeth
are all conical including three small teeth often present on the maxilla.
There are 24-26 anal-fin rays.
Size. This dwarf species grows to about 25 mm TL, but most individu¬
als are smaller.
Pigmentation. The body is mostly transparent but the rear portion and
the tail are bright red or pink and the belly is silvery. The dorsal fin
has a black blotch. Females are not as colorful as males.

425
Figure 163. Megalamphodus cf axelrodi.

Figure 164. Apure drainage distribution of Megalamphodus cf axelrodi.

CUENCA DEL RIO APURE
too
ro

428
Distribution and Natural Hi story
Range. It occurs in Trinidad and in northern South America.
Apure distribution. Map: fig. 164. It has a spotty distribution in
the Apure drainage.
Habitat. Mega!amphodus cf axelrodi is usually found amidst aquatic
vegetation, in quiet sidewaters of slow-moving llanos streams.
Abundance. UNCOMMON.
Number of specimens examined. 83 from 17 collections.
Food. OMNIVORE. It eats microcrustaceans, tiny seeds, and small
insects.
Reproduction. Strategy: rl. It probably spawns throughout the rainy
season. In aquaria, adhesive eggs are scattered over aquatic vegetation and
then abandoned.
Importance. Ornamental.
Metvnnis argenteus Ahl 1923
Palometa Plateada - Silver Dollar
Fig. 165. Map fig. 166. Couplet 17a.
Generic synonymy
Metvnnis Cope 1878:692 (type species: Metvnnis 1 una Cope 1878,
by monotypy).
Mvleocol1 ops Eigenmann 1903:147 (type species: Metvnni s goel di i
Eigenmann 1903, by original designation).
Sealeina Fowler 1906:478 (type species Mvletes 1ippincottianus
Cope 1870, by original designation).
Specific synonymy
Metvnnis argenteus Ahl 1923:24 (type local ity: Río Tapajós [tributary of the
lower R. Amazonas]), seen; Mago L. 1970:71; Géry 1977:270, 271 (key).

429
Figure 165. Metvnnis argenteus.

430
Comments. The genus Metynnis, as is true of most characid genera, is
in need of revision. There are several names available for the Venezuelan
species, but most were described from geographic localities far from Vene¬
zuela.
Etymology. METYNNIS = with a plow, referring to the keeled belly;
ARGENTEUS = silvery.
Description
Illustrations. Fig. 165; Géry 1977:280.
Diagnosis. This species is distinguished from other species of the
genus by the following combination of characters: gill rakers on the lower
limb of first arch 17-23; supraoccipital process relatively short, and
measuring more than 2.6 times in the distance from the base of the occiput
to the dorsal-fin origin; adipose fin not extremely long and short, the
distance from the rear margin of the dorsal-fin base to the adipose fin
measuring 1.5 times in the length of the adipose-fin base. They also might
be confused with small Mvlossoma or Myleus, but members of those genera have
a much shorter adipose-fin base (the adipose-fin base length is less than
the distance from the posterior edge of the dorsal-fin base to the origin of
the adipose fin (fig. 33).
Size. It grows to about 150 mm SL.
Morphology. The body is discoid.
Counts. DR ii16-18; AR 34; abdominal scutes 34.
Measurements. GBD 80-88 % of SL (N=8).
Pigmentation. The body is silvery, with a purple to orange sheen. The
dorsal fin usually has several irregular black spots. The caudal and anal
fins are faint orange to pink.

Figure 166. Apure drainage distribution of Metynnis argenteus.

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Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map: fig 166. This species is known from a few
localities in northern Apure state, and the upper Aguaro River system.
Habitat. It is found both in running and quiet waters, usually asso¬
ciated with aquatic vegetation.
Abundance. UNCOMMON. The relative scarcity of this species may be
more a reflection of difficulties in collecting, rather than true rarity
Number of specimens examined. 178 from 24 collections.
Food. OMNIVORE. It is mostly vegetarian, consuming seeds, flowers,
fruits, aquatic plants.
Reproduction. Strategy: r2. It is probably synchronized with the
onset of the rains in May to June. A pair that spawned in aquaria released
eggs early in the day, before dawn. The pair scattered the eggs at random
after violent side-by-side quivering. It probably spawns over aquatic
vegetation in nature.
Migrations. These fish live much longer than a year, and so probably
make biannual migrations to stay in favorable habitat with the alternation
of the wet and dry seasons.
Importance. It is a highly prized ornamental species.
Metvnnis hypsauchen (Müller & Troschel) 1844
Palometa - Silver Dollar
Fig. 167. Map: fig. 168. Couplet 17b.
Specific synonymy
Mvletes hypsauchen: Muller & Troschel 1844:97 (type locality: British
Guiana), seen.

434
Figure 167. Metvnnis hypsauchen.

435
Metvnnis hypsauchen Eigenmann & Ogle 1907:35 (South America); Ah1
1923:21 (key); Fowler 1948:394 (synonymy).
Metvnnis orinocensis Steindachner 1908b:359 (type locality: ?Rio
Orinoco).
Metvnnis (Myleocol1 ops) hypsauchen Géry 1972:190.
Comments. According to Géry (1977), M. orinocensis Steindachner 1908
is a synonym of this species, but this remains to be proved.
Etymology. HYPS = from the Greek "hypsos" for height, AUCHEN = from
Greek for nape, referring to the deep body.
Description
Illustrations. Fig. 167; Géry 1972:191 fig. 27 (drawing of holotype);
1977:277 (photo).
Diagnosis. This species is distinguished from other members of the
genus by the following combination of characters: gill rakers on lower 1 imb
of first arch about 35; supraoccipital process very long, and measuring
less than 2.4 times in the distance from the base of the occiput to the
dorsal-fin origin; the adipose fin is extremely long and low, the distance
from the rear margin of the dorsal-fin base to the adipose fin measuring
more than 1.5 times in the length of the adipose-fin base. See diagnosis of
M. argenteus to distinguish this species from those of other similar
genera.
Size. It reaches about 150 mm SL.
Morphology. The body is discoid.
Counts. DR ii16-18; A 39-44; abdominal scutes 26-35.
Measurements. GBD 79% SL; HL 3.75 in SL.
Pigmentation. The body is silvery with a few dots present in the
dorsal fin. During the breeding season adults take on an orange sheen.

Figure 168. Apure drainage distribution of Metynnis hypsauchen.

—I
CUENCA OEL RIO APURE
LA UNIVERSIDAD O'lE SlEMBF
UNIVERSIDAD NACIONAl EXPERIMENTAL
DE LOS LLANOS OCCIDENTALES
"EZEOUIEL ZAMORA"
VICE-RECTORADO tN PORTUGUESA
A • •<

438
Distribution and Natural History
Range. It is found in the Amazon and Orinoco basins, and Guyana.
Apure distribution. Map: fig. 168. It is restricted to the Aguaro
River system.
Habitat. It has been found in lentic and slower-moving sectors of
lotic systems.
Abundance. RARE.
Number of specimens examined. 8 from 3 collections.
Food. OMNIVORE. It is mostly vegetarian, consuming seeds, fruits,
flowers, aquatic plants.
Reproduction. Strategy: r2. It is probably synchronized with the
onset of the rains in May to June.
Migrations. These fish live much longer than a year, and so probably
make biannual migrations to stay in favorable habitat with the alternation
of the wet and dry seasons.
Importance. Ornamental.
Metvnnis 1 una Cope 1878
Palometa - Bleeding Heart Silver Dollar
Fig. 169. Map: fig. 170. Couplet 16a.
Specific synonymy
Metvnnis luna Cope 1878: 692 (type locality: Peruvian Amazon), not seen;
Ahl 1923:20 (key).
Comments. This is the type species of the genus. It is the only
Metvnnis with such a high number of gill rakers. The specific identifica¬
tion of this and other Metvnnis species is difficult, and uncertain due to
the plethora of synonyms in the literature, which are mostly based on incom¬
plete original descriptions.

439
Figure 169. Metvnnis luna.

440
Etymology. LUNA = moon shaped.
Description
Illustrations. Fig. 169; Fowler 1950:396 fig. 440; Géry 1977:276;
Román 1985:117.
Diagnosis. This is the only Metynnis species with more than 50 total
gill rakers on the outer arch. See diagnosis of M. argenteus for distin¬
guishing characters of this species from those of other similar genera.
Size. It grows to about 150 mm SL.
Morphology. The discoid body comes to a "peak" with the apex at the
dorsal-fin origin.
Counts. DR 18; AR 39; abdominal scutes 25.
Measurements. BD 80% SL; GBD 11/12 SL.
Pigmentation. This fish is silvery in life, with a red spot in the
humeral area just behind the opercle.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map: fig. 170.
Habitat. It has been found in lentic biotopes and slower-moving
streams in the lower llanos, where it is usually associated with aquatic
vegetation.
Abundance. UNCOMMON.
Number of specimens examined. 89 from 15 collections.
Food. OMNIVORE. It is mostly vegetarian, consuming seeds, fruit,
flowers, and aquatic plants.
Reproduction. Strategy: r2. It is probably synchronized with the
onset of the rains in May to June.

Figure 170. Apure drainage distribution of Metynnis luna.

CUENCA DEL RIO APURE
nut
â– P*
ro

443
Migrations. These fish live much longer than a year, and so probably
make biannual migrations to stay in favorable habitat with the alterna¬
tion of the wet and dry seasons.
Importance. Ornamental.
Microschemobrvcon casiauiare Bohlke 1953
Sardinita de Casiquiare - Casiquiare Tetra
Fig. 48 (caudal pattern) & 171. Map: 172. Couplet 67a.
Generic synonymy
Microschemobrvcon Eigenmann 1915a:56 (type species: Microschemobrycon
guaporensis Eigenmann 1915a, by original designation).
Specific synonymy
Microschemobrycon casiquiare Bohlke 1953b:841 (type locality: Sao
Gabriel Rapids of the Rio Negro, Brazil, shortly downriver from 0°
08’ 03" South Latitude, 67° 03’ 28" West Longitude, where the village
of Sao Gabriel is situated), seen; Mago L. 1970:71; Géry 1977:590-1,
598 (key).
Types. Holotype: CAS 17497. Paratypes: CAS 17498(8); 17499(3);
17500(11); 17501(2); 17502(1); 17503(1); 17504(11); 17505(2); 17506(1);
17507(8); 17508(4); 17509(6); 17510(1); 17511(29); 17512(2); 20689(10);
20690(5).
Etymology. MICRO = small, SCHEM0 = Greek for arrangement or plan,
thus small of stature (Eigenmann 1919); BRYCON = a genus of characid; CASI¬
QUIARE = after the Casiquiare Canal in Venezuela, where the fish was origi¬
nally found.
Description
Illustrations. Fig. 48 (caudal pattern) & 171; Bohlke 1953d:pi.
20).

444
Figure 171. Microschemobrvcon casiauiare.

445
Diagnosis. The humeral spot is absent. There is a dark crescent of
embedded pigment at the base of each caudal -fin lobe (fig. 48). The premax¬
illary teeth are arranged in a single row.
Size. This is a small characid, rarely exceeding 40 mm SL.
Morphology. The body is elongate and slender, cigar-shaped, not
greatly compressed. The dorsal -fin origin is about equidistant from the tip
of the snout and the mid-caudal base. The lateral line is incomplete, with
18-23 pored scales.
Counts. DR ii8-ii9; AR i ii15-i ii18; PR ilOi; VR i7; LS 26 (32-33 in
type series); TS 4 above LL, 2 below. Premaxilla with 12-13 teeth in single
row, maxilla with 5, dentary with 11 large tricuspid teeth followed by 7
smaller conical teeth; GR 9-11.
Measurements. HL 3.4-4.0 in SL, GBD 3.7-4.6 in SL.
Pigmentation. The body is tan and the fins are clear.
Distribution and Natural History
Range. It occurs in the Rio Negro drainage (Amazon), the Casiquiare
Canal, and the upper Orinoco Basin in Colombia and Venezuela.
Apure distribution. Map: 172. It is known only from the Aguaro River
system.
Habitat. It is found in blackwater streams and morichales, usually
near aquatic vegetation.
Abundance. UNCOMMON.
Number of specimens examined. 41 from 6 collections.
Food. MICROCARNIVORE. It probably feeds on microcrustaceans, and
small insects.
Reproduction. Strategy: rl. It probably spawns throughout the
rainy season.

Figure 172. Apure drainaqe distribution of Microschemobrycon casiquiare.

I
I
VICt RECTURADU tN POHTUGUESA
4 • •(
CUENCA OEL RIO APURE
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Moenkhausia chrvsargvrea (Giinther) 1864
Palometica Aro de Oro - Golden-Circle Tetra
Fig. 173. Map: fig. 174. Couplet 94a.
448
Generic synonymy
Moenkhausia Eigenmann 1903:145 (type species: Tetragonopterus xinouensis
Steindachner 1882:178, by original designation).
Specific synonymy
Tetragonopterus chrvsargyrea Giinther 1864:328 (type locality:
Essequibo, British Guiana), seen.
Moenkhausia chrysargyrea Eigenmann 1910:437 (Brazil); 1912:323 (key,
diagnosis, synonymy); 1917:75 (diagnosis).
Etymology. MOENKHAUSIA = to honor Professor W. J. Moenkhaus, of
Indiana University (where Dr. Eigenmann worked for many years) and who
for a time collaborated with the Museu Paulista, in Sao Paulo, Brazil;
CHRYSAR = gold, from the Latin "chrysos," and GYREUS = circle, from the
Latin "gyrus." The significance of this remains obscure. While there
is no golden circle in preserved material, I suspect that the pale halo
around the first humeral spot is probably golden in life.
Description
Illustrations. Fig. 173; Eigenmann 1917:pi. 6, fig. 2.
Diagnosis. The first humeral spot in this species is dark black,
round in shape, and surrounded by a pale area. The second humeral
spot, if present, is faint and vertically elongated. The caudal fin
lacks black markings of any kind. The body is very deep, with its
maximum depth about 2.0-2.1 in the SL.
Size. Although reported to reach 100 mm, most individuals are
between 30-50 mm SL.

449
Figure 173. Moenkhausia chrvsarqvrea.

Figure 174. Apure drainage distribution of Moenkhausia chrysargyrea.

CUENCA DEL RIO APURE
OCAL A •»»»<:
100
-P*
en

452
Morphology. The body is deep, highly compressed and subrhomboi-
dal. The interorbital width is about equal to the eye diameter. The
scales are deeply imbricate, and the lateral line is complete.
Counts. DR 11; AR 27-30; LLS 33-35; PDS 10; premaxilla with 4-5
teeth in outer row, 5-6 pentacuspid teeth in inner; maxilla with 1-2.
GR 14.
Measurements. HL 3.5-4 in SL; GBD 2 (in adults) to 2.4 in young.
Pigmentation. This fish is iridescent silvery, with a deep-
lying, well-defined, horizontally oval, circular, or rhomboidal black
spot over the space between the fifth to the eighth scale of the later¬
al line. A second, faint humeral spot is sometimes present but there is
no caudal spot.
Distribution and Natural Hi story
Range. It occurs in Guyana and Venezuela.
Apure distribution. Map: fig. 174. Two disjunct population have
been found. One in Barinas state near Socopó, the other in Guárico, in
the Aguaro River system.
Habitat. It inhabits quiet llaneran waters.
Abundance. UNCOMMON.
Number of specimens examined. 38 from 6 collections.
Food. OMNIVORE.
Reproduction. Strategy: rl. It probably spawns throughout the
rainy season.
Importance. Ornamental.
Moenkhausia copei (Steindachner) 1882
Bobita, Sardinita - Cope’s Tetra
Fig. 175. Map: fig. 176. Couplet 81a.

453
Figure 175. Moenkhausia copei.

454
Specific synonymy
Tetragonopterus copei Steindachner 1882:179 (type locality: Santarem),
seen; 1883:35.
Moenkhausia copei Eigenmann 1910:439; 1917:pi. 101; 1918:pi.9 (key,
redescription); Mago L. 1970:71; Géry 1977:424, 442, 446.
Comments. Two very similar sympatric species, M. copei and M. col -
lettii, were described by Steindachner from Brazil, and included in the
faunal work of Guyana by Eigenmann (1912). Eigenmann later described the
very similar (if indeed not identical) Hemigrammus barrigonae from Barrigo¬
na, Colombia, in the Andean foothills drained by tributaries of the Meta
River. It seems that since then the three species have often been confused.
The most common form in the Apure drainage is elongate and slender, and has
black pigment of the lateral stripe not extending onto the base of the
caudal rays. I am calling this species M. copei, in keeping with recent
work publ ished (Machado A. 1987). The other deeper-bodied species has black
pigment on the base of the central caudal rays, and is usually much less
common when they occur together. This species is either M. collettii (type
locality middle Amazon, Brazil) or H. barrigonae. It is interesting to note
that both of the latter species are reported to have variable development of
the lateral line, which in some individuals is incomplete, and in others is
nearly complete. Since completeness of the lateral line has been a primary
feature used in defining a number of genera of small characoid fishes, it
becomes clear that this character needs to be reevaluated.
Although there are some differences in the counts of anal -fin rays and
gill rakers between Eigenmann’s different works on these characids and Apure
drainage specimens, they are slight and the type series are small. I have
opted to use the name H. barrigonae for the deeper bodied species, since the
sometimes incomplete lateral line would appear to fit a Hemigrammus

455
better than a Moenkhausia (at least as currently understood) and because it
is described from the same basin. A detailed comparison of material from
all along the Andean Piedmont and into Guyana and Brazil is needed in order
to determine if these taxa represent a cline or a complex of species.
Etymology. COPEI = to honor Edward Drinker Cope, North American
ichthyologist and paleontologist.
Description
Illustrations. Fig. 175.
Diagnosis. This species is distinguished from other members of the
genus by the pigmentation pattern (see Pigmentation below). It is very
similar to Hemigrammus barrigonae. but is more elongate and has fewer seales
between the lateral line and the dorsal-fin origin (5 vs 6-7).
Size. It can grow to about 60 mm SL, but most individuals are between
40-50.
Morphology. The lateral line is variable but usually complete.
Counts. DR 11; AR 19-20; LLS 31-34; TS 10-13; PDS 10-12; GR 10. 4-5
teeth on outer premaxillary row, 5 in inner.
Measurements. GBD 31-38% SL; HL 28% SL; PDL 51% SL.
Pigmentation. The characteristic marking of this species is the thin
black line above the anal-fin base. There is usually no caudal spot as such,
and the lateral stripe ends on the caudal peduncle and does not extend onto
the central caudal-fin rays. The humeral spot is well defined. The
unpaired fins are tinged with red in life.
Distribution and Natural History
Range. It occurs in the Orinoco and Amazon basins, and in Guyana.
Apure distribution. Map: fig. 176. It is known from isol ated popul a-
tions in Barinas state near Socopó, in northern Apure state west of Mante-
cal, and in the Aguaro River drainage.

Figure 176. Apure drainage distribution of Moenkhausia copei.

CUENCA OEL RIO APURE
UNIVERSIDAD NACIONAL EXPERIMENTAL
DE LOS LLANOS OCCIDENTALES
"EZEOUIEL ZAMORA"
VICE-RECTORADO EN PORTUGUESA
herida 4
457

458
Habitat. It inhabits blackwater streams.
Abundance. COMMON.
Number of specimens examined. 1,378 from 42 collections.
Food. OMNIVORE. It eats microcrustaceans, insects, and seeds.
Reproduction. Strategy: rl. It probably spawns throughout the
rainy season.
Importance. Ornamental.
Moenkhausia dichroura (Kner) 1859
Sardinita Tijeras - Scissor-Tail Tetra
Figs. 60 (caudal pattern) & 177. Map: fig. 178. Couplet 95a.
Specific synonymy
Tetragonopterus dichrourus Kner 1859:80 (type locality: Río Guaporé,
Caicara, Paraguay), seen.
Salmo 1ambari Natterer in Kner 1859:177 (name in text).
Poecilurichthys dichrourus Eigenmann & Kennedy 1903:522.
Moenkhausia dichrourus Eigenmann 1907:138; 1910:438.
Moenkhausia dichroura Bertoni 1914:12 (Paraguay); Eigenmann 1912:326;
1917:95; Mago L. 1970:71; Géry 1977:429, 442, 446, 450, 502 (key).
Moenkhausia dicroura (error) Schindler 1939:288 (Paraguay).
Moenkhausia intermedia Román 1985:429 (Venez.).
Etymology. DI = two, CHROURA = colors, probably referring to the
distinctive tail with black and white.
Description
Illustrations. Fig. 60 (caudal fin) & 177; Axelrod et al. 1971:F-
406.00; Géry 1977:429; Román 1985:175, (as M. intermedia).
Diagnosis. The caudal pigmentation pattern (fig. 60) is diagnostic
for this species. It could be confused with Hemigrammus marginatus,
but the latter has an incomplete lateral line.

459
Figure 177. Moenkhausia dichroura.

Figure 178. Apure drainage distribution of Moenkhausia dichroura.


462
Size. The reach about 75 mm TL, but are more commonly taken at around
40 mm.
Morphology. The body is elongate but not spindle shaped, and slight¬
ly compressed. The lateral line is complete.
Counts. DR 11; AR 28; LLS 34.
Pigmentation. The body is silvery and the fins are clear except for
the caudal, which is yellowish and has each milk-white tipped lobe crossed
with a black bar (fig. 60).
Distribution and Natural Hi story
Range. It is widespread from Venezuela to Paraguay.
Apure distribution. Map: fig. 178. This species is known from both
llaneran and piedmont streams.
Habitat. It occurs in lowland llanos.
Abundance. COMMON.
Number of specimens examined. 2,210 from 118 collections.
Food. OMNIVORE. It feeds on seeds and insects.
Reproduction. Strategy: rl. It probably spawns throughout the
rainy season.
Importance. Ornamental, but quite sensitive to handling.
Moenkhausia lepidura complex (Kner) 1859
Sardinita - Tetra
Fig. 60 (caudal pattern) & 179. Map: fig. 180. Couplets 80a & 95b.
Specific synonymy
Tetragonopterus lepidurus Kner 1859:40 tab. 8, fig. 20 (type locality:
Río Guaporé), seen; Moenkhausia lepidurus Eigenmann 1910:438.
Moenkhausia lepidura Eigenmann 1912:326; 1917:98 (key, description,
list of subspecies); Géry 1977: 428, 432, 442, 451 (key).

463
Figure 179. Moenkhausia 1 epidura complex.

464
Comments. This species is part of a complex of species that all have
similar color patterns. Several subspecies have been named, but confusion
still exists because of a lack of adequate comparative material and brief,
incomplete descriptions.
Etymology. LEPIDU = from the Latin "lepidus" for pleasant, neat or
pretty; URA = from the Greek "oura" for tail.
Description
Illustrations. Fig. 60 (caudal pattern) & 179; Eigenmann 1917:pi.
8, fig. 2; Géry 1977:428.
Diagnosis. This species could be confused with M. dichroura or
Hemiorammus marginatus. It differs in having only the upper caudal-fin
lobe darkly pigmented, instead of both. It is also similar to
Brvconops melanurus, but is distinguished in having a shorter maxilla
and fewer lateral-line scales (about 35 vs 43-45).
Size. It can reach almost 110 mm SL, but most individuals are small¬
er, 40-80 mm.
Morphology. The body is long and slender but not particularly com¬
pressed.
Counts. DR 11; LS about 35; AR about 29; GR 9.
Measurements. HL 3.75-4.4 in SL; GBD 3-3.5 in SL.
Pigmentation. The caudal fin has a dark edge on the upper lobe,
and is orange near its base.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins, and in Guyana.
Apure distribution. Map: fig. 180. It occurs only in the llanos.
Habitat. M. lepidura lives in streams of the lowland llanos.
Abundance. COMMON.

Figure 180. Apure drainage distribution of Moenkhausia lepidura complex.


467
Number of specimens examined. 722 from 44 collections.
Food. OMNIVORE.
Reproduction. Strategy: rl. It probably spawns throughout the
rainy season.
Importance. Ornamental.
Myleus cf pacu (Schomburgk) 1841
Palometa - Pacu, Silver Dollar
Fig. 181. Map: fig. 182. Couplet (not in key).
Generic synonymy
Myleus Müller & Troschel 1844:98 (type species: Myleus setiqer Müller
& Troschel 1844, (= Myleus pacu Schomburgk 1841), by subsequent
designation of Eigenmann 1910:443).
Tometes Valenciennes in Cuvier & Valenciennes 1849:225 (type species:
Tometes trilobatus Valenciennes 1849, (= Myleus pacu Schomburgk 1841),
by subsequent designation of Jordan 1919:242).
Myloplus Gill 1896:214 (type species: Myletes asterias Müller &
Troschel, by subsequent designation of Jordan (1920), proposed as
substitute for Myletes of Müller & Troschel).
Orthomyleus Eigenmann 1903:148 (type species: Myletes el 1 ipticus
Günther, by original designation).
Paramvloplus Norman 1929:828 (type species: Paramvloplus ternetzi Norman
1929, by original designation).
Prosomvleus Géry 1972:182 (type species: Myletes rhomboidalis Cuvier
1818, by original designation).
Specific synonymy
Myletes pacu Schomburgk 1841:236, pis. 20-21 (type locality: Guyana),
seen.

468
Figure 181. Myleus cf pacu.

469
Mvleus setiger Müller & Troschel 1844:98 (Guyana).
Mvletes setiger Gunther 1864:378 (Guyana).
Myletes schomburqki (non Jardine i_n Schomburgk) Muller & Troschel
1844:98.
?Myl etes di vari catus Valenciennes i_n Cuvier & Valenciennes 1849:215
(type locality: Essequibo).
Tometes tri1obatus Valenciennes jn Cuvier & Valenciennes 1849:27 (type
locality: Cayenne).
Tometes unilobatus Valenciennes jn Cuvier & Valenciennes 1849:27 (type
locality: Cayenne).
Myletes rhomboidalis (non Cuvier) Pellegrin 1908:28.
Mvleus pacu Eigenmann 1912:393 (Guyana).
Mvleus (Mvleus) pacu Gérv 1972:178 (diagnosis, synonymy); 1977:266
(diagnosis).
Comments. Because only one juvenile specimen, tentatively identified
here as Mvleus pacu (a species hitherto known only from Guyana), was cap¬
tured, this species was not included in the key. That specimen would key
out with the Mvlossoma species, but has more ventral serrae (28-46 vs 10-
22). The similar Mvleus micans is present in morichal habitats in other
parts of Venezuela (Machado Allison, pers. com.), but differs from M. pacu
in various meristic characters (Géry 1977). Since even the identification
of adults is difficult in this group, this name is only tentative, pending
collection of larger specimens from the Aguaro River.
Etymology. MYLEUS = like a mill stone; PACU = is the common name
of this species in Guyana.
Description
Illustrations. Fig. 181; Myers 1972:57; Géry 1977:264.
Diagnosis. The two species of Myleus known from the Apure drainage
are easily distinguished by the dorsal-fin ray count, about i ii20 in

470
M. cf pacu and ÜÍ26-29 in M. rubripinnis. This species is further
distinguished by the following combination of characters: front teeth
compressed, close-set, sharp cutting incisors, appressed against the
inner molariform teeth (Géry 1977); anal-fin rays i i i31.
Size. It reaches at least 600 mm SL (Géry 1977).
Morphology. The body is discoid.
Counts. There are 28-47 ventral serrae (the number increases with
age) (Géry 1977).
Pigmentation. The juvenile specimen collected during this study is
gray dorsally, whitish to silvery ventrally, and with faint irregular darker
vertical bars on the upper sides. Adults are reported to be plain as
adults, though they have large red spots on the sides during the breeding
season (Géry 1977).
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins, and in Guyana.
Apure distribution. Map: fig. 182. This species is known from
one locality in the Aguaro River system.
Habitat. The specimen collected was taken from a blackwater stream
with abundant aquatic vegetation, in the low llanos.
Abundance. RARE.
Number of specimens examined. 1 from 1 collection.
Food. OMNIVORE. It is mostly vegetarian.
Reproduction. Strategy: r2. It is probably synchronized with the
onset of the rains in May to June.
Migrations. It probably migrates at the end of rainy season to
avoid drought conditions.
Importance. Ornamental.

Figure 182. Apure drainage distribution of My1eus cf pacu.

Etc I 2 000000
472

473
Mvleus rubripinnis (Müller & Troschel) 1844
Palometa Gancho Rojo - Redhook Metynnis
Fig. 183. Map: fig. 184. Couplet 18a.
Specific synonymy
Myletes rubripinnis Müller & Troschel 1845:97 (type locality: Guiana).
Myloplus rubripinnis Eigenmann 1912:391 (Guyana); Fowler 1950:404
(synonymy, range); Mago L. 1970:72.
Myleus rubripinnis Eigenmann 1910:443.
Myleus (Myloplus) rubripinnis Géry 1972:162 (diagnosis, synonymy);
1977:262 (key, subspecies).
Etymology. RUBRI= red, PINNIS = fins.
Description
Illustrations. Fig. 183; Fowler 1951:405; Géry 1977:263; Axelrod
et al. 1971:F-420.12.
Diagnosis. The high number of dorsal-fin rays (29 or more), and
the elongate, hooked anal-fin of the male distinguish this species from
the all other discoid characids in the Apure drainage.
Size. It matures at about 90-120 mm (Axelrod et al. 1971), but can
get somewhat larger.
Morphology. The body is discoid. A distinctive red hooked anal
fin is present in adult males.
Counts. DR iii26 to i i i 29; AR ÍÜ32-36.
Pigmentation. The body is silvery and the anal fin red. Breeding
males have brilliant patches of orange, red or yellow on the sides.
Distribution and Natural Hi story
Range. It occurs in the Orinoco and Amazon basins.
Apure distribution. Map: fig. 184. It is known from Caño Caicara
in northern Apure state west of Mantecal, and the Aguaro River system.

474
Figure 183. My1eus rubripinnis. A mature male is shown below.

Figure 184. Apure drainage distribution of Myleus rubripinnis.

476

477
Habitat. It lives in the low llanos, in blackwater streams, with
abundant aquatic vegetation.
Abundance. UNCOMMON.
Number of specimens examined. 13 from 10 collections.
Food. OMNIVORE. It is mostly herbivorous.
Reproduction. Strategy: r2. It is probably synchronized with the
onset of the rains in May to June. Males are distinguished by having the
central rays of the anal fin elongated and hooked. In captivity, 300-500
eggs per females are scattered randomly about the tank. The embryos hatch
in about 50-60 hours, absorb the yolk-sac over a period of 3-7 more days,
and then begin to feed on plankton (Axelrod et al. 1971).
Migrations. Since this is a fairly large fish, dry-season migrations
to avoid drought conditions may be expected, but have not yet been document¬
ed in the Apure drainage.
Importance. Ornamental.
Mvlossoma aureum (Spix) 1829
Palometa Dorada - Golden Palometa
Fig. 185. Map: None. Couplet 31b.
Generic synonymy
Mylossoma Eigenmann & Kennedy 1903:530 (type species: Myletes albiscopus
Cope, by monotypy).
Mylosoma Eigenmann 1915:261 (alternative spelling).
Starksina Fowler 1906:476 (type species: Mvletes herniarius Cope,
by monotypy).
Specific synonymy
Myletes aureus Spix in Agassiz 1829:74 (type locality: Equatorial rivers
of Brazil), seen.

478
Figure 185. Mylossoma aureum.

479
Tetragonopterus aureus Spix in Agassiz 1829:74 pi. 31.
Mvlosoma aureum Eigenmann 1910:444.
Mvlosoma aureus Eigenmann 1915:265 (Brazil).
Mylossoma aureus Pearson 1925:48 (Bolivia).
Mvlossoma aureum Norman 1928:812 (Orinoco, Amazon); Eigenmann and Allen
1942:249 (Orinoco to Ucayali and La Plata); Schultz 1944b:258; Fowler
1945:168; Mago L. 1970:72; Géry 1977:258-9, 261.
Myletes herniarius Cope 1872:268, pi. 12 fig. 3 (type locality: Rio
Ambyiacu, Peru).
Starksina herniarius Fowler 1907:476, fig. 56 (types).
Colossoma herniarium Eigenmann 1910:444.
Mylossoma herniarium Ahl 1922: non-paginated.
Myletes albiscopus (not of Cope) Boulenger 1898:426 (Río Jurúa) (part).
?Metynnis unimaculatus Steindachner 1908:326 (type locality: lake near
Rio Medonho, trib of R. Parnaiba, north of Santa Filomena).
Mylossoma duriventris (not of Cuvier) Ahl 1922 (figures only).
Comments. This is something of a "mystery fish," since it is always
cited, but few specimens in collections are identified as such. I suspect
that it goes unnoticed among the more common M. duriventris.
Etymology. MYLOS = millstone, SOMA = body; AUREUM = golden.
Description
Illustrations. Fig. 185; Fowler 1951:407 fig. 451 (type of Myletes
herniarius Cope; Myers 1972:41; Román 1985:115.
Diagnosis. Mylossoma species differ from Colossoma in possessing a
heavily scaled anal fin (vs scaled only at base), in having only four teeth
on each side of the lower jaw (vs six or more), and in having more than 35
anal-fin rays (vs less than 30). They are distinguished from the other

480
large discoid serrasalmine genera (for example Mvleus and Metvnnis) in
lacking the predorsal "spine" that is embedded in the flesh just ante¬
rior to the dorsal-fin origin in those genera.
Mvlossoma aureum differs from Mvlossoma duriventris in having
fewer serrae behind the pelvic fins (10-16 vs 18-22), with the last one
not close to the first anal-fin ray; in having only 28-34 branched
anal-fin rays (vs about 37), and in having a relatively shorter adi¬
pose-fin base measuring 3.75-4.25 times in the dorsal-fin base length
versus 2.6-3.66 (Géry 1977).
Size. It grows to about 300 mm SL.
Morphology. It is plate-shaped, very deep-bodied and compressed.
Counts. DR Ü13-15.
Measurements. GBD 70% SL.
Pigmentation. The body is silvery to golden.
Distribution and Natural History
Range. It occurs from the Orinoco in Venezuela to the Ucayali and
La PI ata in Brazil.
Apure distribution. Map: No specimens were collected during this
study, but many have been collected by fishery biologist Otto Castillo
from around San Fernando de Apure and (pers. com.).
Habitat. It inhabits large river channels and llaneran creeks with
abundant overhanging shore vegetation. It often seeks refuge in submerged
brush piles near shore.
Abundance. RARE. The relative scarcity of this species may be more a
reflection of its ability to avoid capture with the equipment used, rather
than true rarity. It is not as common as M. duriventris, and the two are
difficult to distinguish without careful examination.
Number of specimens examined. 0.

481
Food. OMNIVORE. It is mainly vegetarian, taking both leafy
aquatic macrophytes as well as fruits and seeds.
Reproduction. Strategy: r2. It is probably synchronized with the
onset of the rains in May to June.
Migrations. It probably makes biannual migrations, upstream at the
beginning of the dry season, and back downstream with the onset of the
rains.
Importance. It is fished commercially and by sport fishermen along
with the more common M. duriventris.
Mvlossoma duriventris (Cuvier) 1818
Palometa - Silver Dollar
Fig. 186. Map: fig. 187. Couplet 31a.
Specific synonymy
Mvletes duriventris Cuvier 1818:451:pi. 22, fig. 2 (type locality:
Brazil), seen; ?Peters 1877:473 (Calabozo, Venez.).
Mvletes duriventre Steindachner 1917:pi. 6 figs 1-3.
Mvletes (Mvlosoma) duriventris Steindachner 1917:55 (Rio Negro).
Mvlossoma duriventris Ah1 1922: non-paginated (2 figs.); Schultz
1944b:258; Mago L. 1970:72; Novoa et al. 1982:271 (biology).
Mvlossoma duriventre Campos 1944:207 (Rio Miranda).
Mvletes albiscopus Cope 1872:267, pi. 12 fig. 1 (type locality: Rio Ambyia-
cu).
Mvlossoma albiscopus Eigenmann & Kennedy 1903:530 (Paraguay).
Mvlosoma albiscopus Eigenmann 1910:444; 1915a:266.
Mvlossoma albiscopum Eigenmann & Allen 1942:250.
Mvlossoma ocel 1 atum Eigenmann 1915a:264 (type locality: Villa Flayes,
Paraguay).

482
Figure 186. Mvlossoma duriventris.

483
Etymology. DURI= hard, VENTRIS = ventrum, probably referring to the
spiny serrae on the ventral edge of body.
Description
Illustrations. Fig. 186; Géry 1977:261; Machado A. 1982:fig. 6
(juvenile); Novoa et al. 1982:fig. 7.
Diagnosis. See preceding diagnosis for M. aureum.
Size. It can reach 340 mm TL, and 1.1 kg (Novoa et al. 1982).
Morphology. The body is disk-shaped.
Counts. DR Ü13-15; AR about 37.
Pigmentation. The body is silvery, sometimes with a bit of yellow on
the abdomen. The fins are clear.
Pistribution and Natural History
Range. It occurs in the Orinoco, Amazon, and La Plata basins.
Apure distribution. Map: fig. 187. It is common in the larger
creeks and rivers of the llanos.
Habitat. It is found in large creeks and rivers during the dry
season, and in lowland flooded areas (bajios and esteros) during the wet
season.
Abundance. COMMON.
Number of specimens examined. 203 from 49 collections.
Food. HERBIVORE. It feeds on seeds, terrestrial vegetation, aquatic
vegetation, and flowers. Goulding (1980) noted that the teeth of Mvlossoma
spp are heavy but have sharper cusps than the teeth of the larger Colossoma
and Piaractus. and are thus better suited for slicing and cutting, rather
than crushing. It eats smaller seeds and fruits when available, and take
leaves and Ceiba seeds during low-water periods in the Amazon Basin when
food is scarcer.

Figure 187. Apure drainage distribution of Mylossoma duriventris.

-p*
00
c_n

486
Reproduction. Strategy: r2. It is probably synchronized with the
onset of the rains in May to June.
Migrations. At the onset of the dry season it moves out of the
shallow, flooded areas of the lower floodplain (esteros and bajios) into
the main rivers and then travels upstream to more permanent creeks in the
piedmont and high llanos that still have their shore vegetation intact. It
seeks food and refuge from the gallery forest trees and shrubs.
Importance. It is an important food fish, and is caught commercially
and by sport fishermen in huge quantities during their annual upstream
migration at the beginning of the drought.
Paragoniates alburnus Steindachner 1876
Sardina Azul - Blue Tetra
Fig. 188. Map: fig. 189. Couplet 6b.
Generic synonymy
Paragoniates Steindachner 1876:114 (type species: Paragoniates alburnus
Steindachner, by subsequent designation of Eigenmann 1910:441).
Specific synonymy
Paragoniates al burnus Steindachner 1876:1 17, pi.8 fig 3. (type
locality: Rio Amazonas at Teffé), not seen; Bohlke 1958 (description);
Mago L. 1970:72; Géry 1977:341, 346, 347 (key).
Etymology. PAR = like, AGONIATES = Agoniates. a genus of characid;
ALBURNUS = whitish.
Description
Illustrations. Fig. 188; Román 1985:156; Géry 1977:341.
Diagnosis. This species is characterized by the following
combination of characters: dorsal-fin origin behind anal-fin origin;
anal-fin long, with 43-49 rays; premaxilla with seven tricuspid teeth

487
Figure 188. Paraqoniates alburnus.

488
arranged in a single irregular row; maxilla completely toothed with 18-
25 small teeth; lateral line incomplete.
Size. It can grow to at least 100 mm SL.
Morphology. The body is moderately deep but elongate and quite
compressed.
Counts. DR Ü9-Ü10; AR 43-49; PR i 11 -i 12; VR i7; TS 14-16; LS
40-43.
Measurements. HL 3.9-4.2 in SL, GBD 2.7-2.9 in SL EYE 3.0-3.3 in
HL.
Pigmentation. The sides of the body have a bluish iridescent
sheen. The fins are clear to dusky gray. There is a large black
blotch at the base of the caudal fin.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map: fig. 189. It is common in most llaneran
streams and rivers.
Habitat. It is usually taken from near shore, among cover or
aquatic vegetation, in large creeks or rivers with a moderate to strong
current.
Abundance. COMMON.
Number of specimens examined. 222 from 61 collections.
Food. CARNIVORE. It feeds on small fish, insects and other inverte¬
brates.
Reproduction. Strategy: r2. It probably spawns at the beginning of
the rainy season.
Importance. Ornamental.

Figure 189. Apure drainage distribution of Paragoniates alburnus.

-p*
UD
O

ParapriStella georqiae Géry 1963
Sardinita - Tetra
491
Fig. 190. Map: fig. 191. Couplet 34a.
Generic synonymy
Parapristel1 a Géry 1963:25 (type species: Pristel1 a aubynei Eigenmann
1909, by original designation).
Specific synonymy
Parapri stel 1 a georgiae Géry 1963:25 (type local ity: about 200 miles east of
Bogotá, Colombia in the upper Rio Meta drainage); 1977:487, 504.
Etymology. PARA= like, PRISTELLA = Pristella is a similar genus of
Characidae; GEORGIAE = after Dr. Géry’s wife.
Description
Illustrations. Fig. 190; Géry 1977:487 (teeth), 504.
Diagnosis. Parapristel1 a georgiae is distinguished by the following
combination of characters: maxilla with about seven small tricuspid teeth;
caudal-fin base with a triangular spot; anal-fin rays about 16 (fewer than
in similar species of Hemigrammus).
This species is easily confused with Hemigrammus micropterus and
Serrabrvcon magoi which are so similarly marked as to suggest that there
may be some sort of mimicry involved. Closer examination reveals signifi¬
cant differences in the number of anal-fin rays and dentition.
Size. It can grow to about 50 mm SL.
Morphology. The lateral line is incomplete.
Counts. DR 11; AR 16; LLS 30. Premaxillary teeth in two rows, 3-4
conical to tricuspid in the outer, 5 pentacuspid in the inner; maxilla
with 6-8 tricuspid teeth; dentary with 5 pentacuspid teeth in front and 13
smaller tricuspid ones on sides (Géry 1964).

492
Figure 190. ParapriStella qeorgiae.

Figure 191. Apure drainage distribution of Parapristella georgiae.

10*
9*
8*
CUENCA DEL RIO APURE
i 00
(ICALA MAÍlCA
too
-p*
VO

495
Measurements. GBD 3.47-3.84 in SL; HL 3.4 in SL.
Pigmentation. The caudal-fin base is marked with a triangular black
dot, and with a red dot (in life) that is above and slightly anterior to
the black. The dorsum olive-yellow. The dorsal and caudal fins are
yellowish.
Distribution and Natural History
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 191. It has been collected only from
the Aguaro River system.
Habitat. It is found in blackwater streams.
Abundance. UNCOMMON.
Number of specimens examined. 462 from 8 collections.
Food. CARNIVORE. It feeds on microcrustaceans and insects.
Reproduction. Strategy: rl. It probably spawns repeatedly through¬
out the rainy season.
Importance. Ornamental.
Phenacogaster cf megalostictus Eigenmann 1909
Sardinita - Tetra
Fig. 192. Map: fig. 193. Couplet 83a.
Generic synonymy
Phenacogaster Eigenmann 1907:769 (type species: Tetragonopterus
pectinatus Cope 1869-70, by monotypy).
Specific synonymy
Phenacogaster mega!ostictus Eigenmann 1909:28 (type locality:
Tumatumari, British Guiana), seen; 1917:195, fig. 2; 1927:pi. 56, fig.
2; Eigenmann & Myers 1929:448 (key, description); Mago L. 1970:72;
Géry 1977:530, 548 (key).

496
Figure 192. Phenacogaster cf megalostictus.

497
Types. Holotype: originally CM 1056, now at FMNH. Paratypes: CAS-SU
21906 (2); 21907 (1); CAS 48095(3); 62202(10); 62203(2); 62204(3);
62205(3). I also examined paratypes of P. microstictus CAS 62270(3) and
62271(1).
Comments. This species only superficially resembles P. mega!ostictus
and is probably new. The size and intensity of the humeral spot seems to
vary greatly.
Etymology. PHENACO = a cheat, GASTER = belly, referring to the
peculiar (reduced in number) scalation of the preventral area (Eigenmann
1929); MEGAL0 = big, STICTUS = spot, referring to the humeral spot.
Description
Illustrations. Fig. 192; Eigenmann 1929:pi. 56, fig. 2, pi. 95,
fig. 2.
Diagnosis. Phenacogaster cf megalostictus can be recognized by the
following combination of characters; anal-fin origin under that of the
dorsal fin; scales in front of the pelvic fins enlarged, and in two series
that overlap on the midline (midline sometimes with additional small acces¬
sory scales); lateral spot usually present, large and circular; teeth in
premaxillary rows 4-7 in the outer series, 6-10 in the inner.
Size. It has been reported to reach 85 mm SL in Guyanan specimens
(Géry 1977), but the Apure specimens seldom exceed 50.
Morphology. The body is very compressed and elongate, with the
posterior portion of body elongated. The lateral line is complete. The
preventral area is flat, with two series of large scales overlapping along
the middle, and a small auxiliary scale in the angle of each pair of the
overlapping scales, except the two pairs between the pectorals.
Counts. DR 11; AR 33-37; LLS 32-36; GR 9.

Figure 193. Apure drainage distribution of Phenacogaster cf megalostictus.

CUENCA DEL RIO APURE
O
CICALA AAAClCA
-P»
VD

500
Measurements. HL 3.8-4 in SL; GBD 2.7.
Pigmentation. The body is tan, usually with a rounded black humeral
spot (the spot is often diffuse, faded or even absent in some preserved
material).
Distribution and Natural History
Range. It is known from the Essequibo, Orinoco and Amazon basins.
Apure distribution. Map: fig. 193. It has been found only in north¬
ern Apure state and in the Aguaro River system.
Habitat. It inhabits blackwater streams of the low llanos.
Abundance. UNCOMMON.
Number of specimens examined. 183 from 17 collections.
Food. CARNIVORE. It probably feeds on microcrustaceans, small fish,
and insects.
Reproduction. Strategy: rl. It is probably an egg scatterer that
spawns repeatedly throughout the rainy season.
Importance. Ornamental.
Piaractus brachypomus (Cuvier) 1818
Cachama - Giant Pacu
Fig. 39b (operculum) & 194. Map: 195. Couplet 30b.
Generic synonymy
Piaractus Eigenmann 1903: 148 (type species: Myletes brachypomus Cuvier
1818, by original designation).
Regañina Fowler 1906:475 (type species: Myletes bidens Agassiz 1929, by
original designation).
Waiteina Campos 1946:219 (type species: Waiteina fowleri. by original
designation).

501
Figure 194. Piaractus brachypomus.

502
Specific synonymy
Myletes brachypomus Cuvier 1818:452, pi. 22, fig. 1 (type locality:
Brazil), seen; Valenciennes i_n Cuvier & Valenciennes 1849:199
(redescription of holotype).
Mvletes pacu (not of Schomburgk 1841) Humboldt 1817:175, pi. 47, fig. 2
(type locality: Rio Amazonas, Brazil); Salmo pacu Kner 1860:21 (name
in text).
Myletes bidens Agassiz jn Spix & Agassiz 1829:75, pi. 32 (type locality:
rivers of equatorial Brazil);
Colossoma brachypomus Norman 1928:809 (key, synonymy, Amazon); Schultz
1944b (Rio Caripe, near Caripito, Venez.).
Piaractus brachypomus Eigenmann 1903:148; Eigenmann & Allen 1942:247
(range); Machado A. 1982:47 (redescription, range, synonymy).
Regañina bidens Fowler 1906:475.
Colossoma bidens Eigenmann 1910:444; Fowler 1945:167 (synonymy, range);
Waiteina fowleri Campos 1946:219, fig. 2 (type local ity: Río Tapa jos, Brazil.
Colossoma brachypomum Fowler 1950:388 (synonymy, range); Novoa et al.
1982:268 (biology).
Colossoma brachipomus (error) Román 1985:115 (Venez.).
Types. Holotype: MNHN A.8626.
Comments. As the synonymy indicates, the generic placement of this
species has varied a great deal through the years. The most recent treat¬
ment is that of Machado A. (1982), who is revising the subfamily Serra-
salminae. He recognized only two valid species in this genus, including
this species and P. mitrei (Berg) 1895.
Etymology. PIAR = fat, ACTUS = a ray, from the Greek; BRACHY = wide,
P0MUS = cheek.

503
Description
Illustrations. Fig. 39b (opercle) & 195; Géry 1977:252; Machado
A. 1982: (holotype and juvenile phases); Novoa et al. 1982:fig. 5, (as
Colossoma brachypomum); Román 1985:115 (as Colossoma brachipomus).
Diagnosis. There are only two giant characids, this species and
Colossoma macropomum. Piaractus brachypomus has more lateral-line
scales (78-90 vs 67-76); a narrower opercle (fig. 39), and lacks
ossified rays in the adipose fin. The adipose fin gradually degenerates
and is completely lost in the largest individuals (Machado A. 1982).
Size. This is one of the largest characids in Venezuela. A 450 mm
specimen measured fresh weighed 3.1 kg. Novoa et al. (1982) reported a
maximum size of 790 mm TL from the Orinoco delta region. Specimens as
large as 40 kg have been reported by fishermen.
Counts. DR i i-i ii13-15 (15-18); AR iii-iv21-24 (24-28); PR i15-18
(16-19); VR i7; Caudal Rays i-9+8-i; LLS 70-90; TS 22-30 above LL, 22-33
below; GR 26 to 39, increasing with size of specimen.
Measurements. GBD 47-67% SL; HL 29-50% SL (Machado A. 1982).
Pigmentation. Very small juveniles have ovate spots and a small
ocellus on the sides that disappear at about 60 mm SL. A black opercular
spot is present in specimens up to about 100 mm SL. Medium-sized juveniles
mimic Pygocentrus caribe with their bright red abdomen. In adults, the body
is dark gray or brown on the dorsum and sides with a whitish abdomen. A
few individuals have orange to reddish spots on the abdomen. Older speci¬
mens are darker, usually dark brown but sometimes jet-black.
Distribution and Natural History
Range. It occurs from the Orinoco to the La Plata Basin.

504
Apure distribution. Map: fig. 195. It is common in the larger
rivers and streams throughout the llanos.
Habitat. These fish move from habitat to habitat with ease, spending
part of their time in main channels, but also exploring side creeks, and
flooded areas for fruit dropped from trees.
Abundance. COMMON. Though the few collections don’t in themselves
warrant this classification, this species was once one of the most fre¬
quently observed fishes in the local markets. It is seldom taken with
small seines. Its abundance is declining according to commercial catches
recorded in San Fernando de Apure by Otto Castillo (pers. com.).
Number of specimens examined. 31 from 15 collections.
Food. OMNIVORE. This species feeds on fruits, seeds, flowers,
aquatic vegetation, snails, insects and small fishes. Novoa et al. (1982)
reported a seasonal change in its diet. He found it had a mostly vegetar¬
ian diet during the rainy season, and fed principally on a variety of
fruits. In the dry season, it became more carnivorous and ate more insects
and benthic invertebrates.
Reproduction. Strategy: r2. These fishes mature during the late dry
season and reproduce with the first rains. Fecundity is high, with between
200,000 - 1,5000,000 eggs per female (Novoa et al. 1982).
Migrations. It moves upstream in large schools during the dry
season to abandon the drying savanna streams and move into the main
channel and up towards the highlands. It comes back down with the
onset of the rains.
Importance. This is one of the most important food fish in the Apure
drainage. It is highly appreciated by consumers, and usually on of the top
3-4 species in total weight sold in the commercial market.

Figure 195. Apure drainage distribution of Piaractus brachypomus.


507
Poptel1 a orbicularis (Valenciennes) 1849
Palometica - Silver Half-Dollar
Figs. 55 & 196. Map: fig. 197. Couplet 85a.
Generic synonymy
Poptel1 a Eigenmann 1908:106 (type species: Tetraoonopterus lonqipinnis
Popta, by original designation).
Ephippicharax Fowler 1913a:51 (type species: Tetragonopterus compressus
Günther, substitute name for Fowlerina).
Fowlerina (not of Pelseneer 1906 in Mollusca) Eigenmann 1907:771 (type
species: Tetragonopterus compressus Günther, by original designation).
Specific synonymy
Tetragonopterus orbicul aris Valenciennes i_n Cuvier & Valenciennes
1849:138 (type locality: Essequibo River, British Guiana).
Salmo sau-a Natterer in Kner 1859:174 (name in text).
Tetragonopterus compressus Günther 1864:319 (type locality: Essequibo,
Surinam, Marañon).
Gvmnocorvmbus nemopterus Fowler 1914:248 (type locality: Rupununi River,
British Guiana).
Fowlerina orbicularis Eigenmann 1910:441.
Ephippicharax orbicularis 1914:250 (Guyana); Eigenmann & Myers 1929:503
(Rio Meta).
Ephippicharax orbicularis orbicularis Fowler 1950:367.
Poptel!a orbicularis Géry 1977:365-7, 454.
Etymology. POPTELLA = named to honor the naturalist Popta, the
author of the species Tetragonopterus lonqipinnis, type species of the
genus; ORBICULARIS = from the Latin "orbis" for disk, in reference to
the round body shape.

508
Figure 196. Poptella orbicularis.

509
Description
Illustrations. Fig. 55 & 196; Axelrod et al. 1971:F-486.10; Hoedeman
1975:688; Géry 1977:365, 366.
Diagnosis. This species has a predorsal spine embedded in the flesh
just in front of the dorsal-fin origin. No other similar tetra in the
Apure drainage has this structure. It is superficially similar to Cteno-
brvcon spi1urus. but lacks the ctenoid scales typical of that species.
Another look-alike, Gymnocorvmbus thaveri. can be distinguished by having
more lateral-line scales (about 37 vs about 32). Neither of the two species
mentioned have the distinctive color pattern shown for f\ orbicularis in
fig. 55.
Size. It is reported to get quite large, Hoedemann (1975), reaching
up to 120 mm in aquaria, however, the largest specimens taken during this
study didn’t reach half that size.
Morphology. This fish is deep-bodied, discoid, and highly com¬
pressed. A "saddle-shaped" predorsal spine is embedded in the flesh just
anterior to the first dorsal rays.
Counts. AR 35-37; LLS 37.
Pigmentation. The body is silvery with a faint, often double humeral
spot. The first anal-fin rays are black and usually elongated.
Distribution and Natural History
Range. It occurs in the Essequibo, Amazon and Orinoco basins.
Apure distribution. Map: fig. 197. This species is found in the low
11 anos.
Habitat. P. orbicularis is found both in lentic and lotic habitats,
in white and black water, but is more common in the latter.
Abundance. COMMON.

Figure 197. Apure drainage distribution of Poptella orbicularis.

CUENCA DEL RIO APURE
(«CALA MASCA
100

512
Number of specimens examined. 3,662 from 108 collections.
Food. OMNIVORE. It feeds on seeds, spores, and microcrustaceans (K.
Winemiller pers. com.).
Reproduction. Strategy: rl. This species spawns repeatedly through¬
out the rainy season. Females are larger, males have more intense black on
their elongate first anal-fin rays. Eggs and sperm are scattered among
aquatic vegetation and then abandoned (Floedeman 1975).
Importance. This is an important ornamental species.
Pristobrvcon striolatus (Steindachner) 1908
Caribe Palometa - Silver Dollar Piranha
Fig. 37 (caudal fin) & 198. Map: fig. 199. Couplet 28a.
Generic synonymy
Pristobrvcon Eigenmann 1915:245 (type species: Serrasalmo (Pygocentrus)
calmoni Steindachner 1908b, by original designation).
Specific synonymy
Serrasalmo (Pygocentrus) striolatus Steindachner 1908b:360 (type
locality: Río Para).
Serrasalmus striolatus Norman 1928:794.
Serrasalmus (Pristobrvcon) striolatus Géry 1977:279, 281, 283.
Pristobrvcon striolatus Machado A. 1985:19; Nico & Taphorn 1988:311.
Pygocentrus striolatus Nico & Taphorn 1986:33
Comments. Piranha taxonomy is currently being revised by Dr. W. Fink
& A. Machado A. Since available synonymies will undoubtedly undergo major
revisions once their results are known, a greatly reduced list of refer¬
ences is given for this and the other species of piranhas in the Apure
drainage (the genera Pygocentrus, and Serrasalmus).

513
Figure 198. Pristobrvcon striolatus.

514
Etymology. PRISTO = a saw, BRYCON = a genus of characid, hence a
serrate "brycon," referring to the ventral serrae; STRIOLATUS = striated,
from the Latin for furrow, marked with the light and dark bands present
only in juveniles.
Description
Illustrations. Fig. 37 (caudal fin) & 198; Norman 1928:795; Myers
1972:111; Géry 1977:281; Nico & Taphorn 1986:33.
Diagnosis. This is the least pigmented of the piranhas. The body is
silvery with only a few scattered irregular spots and there is no humeral
spot. The caudal peduncle may have a faint black bar that extends out into
the caudal fin along the outer rays (similar to the pigment pattern ob¬
served in S. irritans). There is a thin line of pigment along the base of
the anal fin. The other fins are transparent. Its dentition is also
characteristic. The teeth on the roof of the mouth are squared distally
with rounded bases, and usually number fewer than five on either side. The
last premaxillary tooth is about as wide as the tooth preceding it.
Size. It can reach about 260 mm SL.
Morphology. The body is deep and fairly compressed. The snout is
blunt, with its length shorter than the eye diameter. The dorsal profile
is slightly concave above the eyes. The lower jaw projects slightly beyond
the upper.
Counts. DR Ü13-16; AR i ii29-31; PR 13-14; teeth 6 on premaxilla,
7 on dentary, ventral serrae 20-23 before ventral fin origin + 9 after.
Measurements. GBD 61% SL; HL 3.3-3.4 in SL.
Pigmentation. See diagnosis. This fish is usually plain and silvery
in life, but sometimes shows orange vertical bars on the sides. In preserv¬
ative, the body sometimes shows irregular stripes or bars on the upper

Figure 199. Apure drainage distribution of Pristobrycon striolatus.


517
sides and the operculum has a dark blotch. The base of the caudal fin is
black.
Distribution and Natural History
Range. It occurs in Guyana, and in the Orinoco and Amazon basins.
Apure distribution. Map: fig. 199. This species has been found
only in northern Apure state, west of Mantecal.
Habitat. It lives in blackwater llaneran streams.
Abundance. UNCOMMON.
Number of specimens examined. 40 from 12 collections.
Food. CARNIVORE. Small juveniles start life as pianktivores,
taking copepods, cladocerans, diptera larvae and fish scales. They later
switch to fish fins scales and flesh, seeds, shrimp, aquatic beetles and
other aquatic invertebrates as main items in their diet (Machado A. &
Garcia 1986; Nico & Taphorn 1986, 1988).
Reproduction. Strategy: r2. It is probably an egg scatterer that
synchronizes reproduction with the onset of the rainy season in May to
June.
Pyqocentrus caribe Valenciennes 1849
Caribe Pechirojo, Colorado or Capaburro - Red-Bellied Piranha
Fig. 200. Map: fig. 201. Couplet 22a.
Generic synonymy
Pyqocentrus Müller & Troschel 1844:94 (type species: Serrasalmo piraya
Cuvier, by subsequent designation of Eigenmann 1910:442).
Gastropristis Eigenmann 1915:238 (type species: Serrasalmo (Pyqocentrus)
ternetzii Steindachner, by original designation).
Rooseveltiella (not of Fox 1914 in Siphonaptera) Eigenmann 1915:240
(type species: Pyqocentrus nattereri Kner, by original designation).

518
Figure 200. Pyqocentrus caribe.

519
Specific synonymy
Serrasalmus caribe Valenciennes 1849:278 in Cuvier & Valenciennes 1849
(type locality: Rio Orinoco, Río Apure, Llanos of Venezuela);
Eigenmann & Eigenmann 1891:60; Norman 1928:803; Schultz 1944b:255.
Serrasal mo (Pygocentrus) notatus Liitken 1874:238 (type locality:
Caracas, Venezuela), not seen.
Serrasalmo nattereri Peters 1877:472 (San Fernando de Apure, Venez.)
Rohl 1942:377 (Orinoco).
Pygocentrus notatus Eigenmann & Eigenmann 1891:60 (Venez.); Taphorn &
Lilyestrom 1984:70 (biology) Machado A. & Garcia 1986:193 (diet); Nico
& Taphorn 1986:24 (diet); Nico & Taphorn 1988:311.
Pygocentrus stiqmatervthraeus Fowler 1911:424 (type locality: La Pedrita
on Caño Uracoa, Venez.).
Rooseveltiella stigmaterythraeus Eigenmann 1915:245.
Serrasalmus notatus Norman 1928:788, fig.5. (as junior synonym of S.
nattereri).
Serrasalmus nattereri Schultz 1944b:254.
Serrasalmus (Taddvella) notatus Géry 1977:291
Comments. According to Drs. W. Fink and A. Machado Allison, who
are currently revising the serrasalmines (pers. com.), the correct name
of the most common piranha in the Apure drainage is P. caribe . This
fish has usually been identified in previous literature as Pygocentrus
notatus.
Etymology. The etymology of this generic name remains unclear,
perhaps PYGO is from the Greek "pygidion" for rump or caudal, or possi¬
bly from the Latin "pygmaeus" for pygmy or small, CENTRUS = central,
middle; CARIBE = the local Venezuelan name for piranha.

520
Description
Illustrations. Fig. 200; as P. notatus: Norman 1928:788 (drawing of
holotype); Mago L. 1978:21; Novoa et al. 1982: fig. 8; Machado A.
1982:fig. 5 (juvenile); Román 1985:103; Nico & Taphorn 1986:24.
Diagnosis. This species is easily recognized in its normal adult
coloration of a steel-blue dorsum and sides, bright scarlet-red belly, and
large jet-black humeral spot. It is further characterized by having asym¬
metrically tri- or pentacuspid teeth that are set so close to one another
as to form a single, jagged cutting edge (so much so that the teeth on
either side of one jaw are replaced as a set, not individually). There are
no teeth on the roof of mouth. The anal-fin rays number 29 or fewer.
Size. Novoa et al. (1982) reported specimens that reached 480 mm TL.
One of the largest specimens from this study, measured 315 mm SL, and
weighed 1 kg.
Morphology. The body is thick, heavy-set and somewhat discoid. The
snout is quite blunt and short, the lower jaw is massive and projects far
beyond the upper.
Counts. DR Ü15-17; AR 26-29.
Pigmentation. The dorsum is steel-blue, olive-green or gray. The
sides are similarly colored but more silvery. The abdomen, and the pector¬
al, anal and pelvic fins are bright scarlet red. A distinctive, large
black humeral blotch is present just behind the opercle. Adults in breed¬
ing color are black with purple and gold flecks on the dorsum and sides.
Distribution and Natural Hi story
Range. It is known from the Orinoco Basin.
Apure distribution. Map: fig. 201. It is very common in lowland
llanos, but extends up to the piedmont.

Figure 201. Apure drainage distribution of Pygocentrus caribe.

522

523
Habitat. It lives in all types of water both lentic and lotic, in
the llanos and piedmont.
Abundance. COMMON. This is the most widespread species of piranha
in the Apure drainage.
Number of specimens examined 1,136 from 174 collections.
Food. CARNIVORE. The smallest juveniles (10-19 mm SL) feed on
microcrustaceans and gradually switch to small aquatic insects. Cladocer-
ans, chironomid larvae, copepods, ostracods and Ephemeroptera make up the
diet at these smallest sizes. Unlike most other piranha species, fish fins
and scales are of minor importance for larger juveniles. As they increase
in size they take larger aquatic insects and then switch to fish flesh as
the major element of their diet for the rest of their lives. Adults feed
mostly on whole small fishes and the fins of larger fishes. They also eat
small vertebrates such as birds, lizards, and mice (K. Winemiller pers.
com.); Machado A. & Garcia 1986; Nico & Taphorn 1986, 1988).
Reproduction. Strategy; r2. The parental protection of the nest
site that is exhibited by this species led us to classify it as a K strate¬
gist in an earlier report (Winemiller & Taphorn 1989). However, its high
fecundity and single reproductive bout per year seem more in accord with r2
strategists. This species’ apparent intermediate nature merely serves to
point out the continuous nature of the life history strategy scheme we
proposed. Most species’ life history strategies are a compromise between
the different selective pressures brought to bear. Sexual maturity is
usually reached by the age of 12 months. Gonads begin to mature in this
species 1-2 months before the onset of the rains. Reproduction is annual
and usually occurs during the first three months of the rainy season, when
most individuals of reproductive age participate in a massive spawning

524
bout. Fecundity is high, between 3048 (Winemiller & Taphorn 1989) in small
females to 20,000 eggs in the largest (González Y. 1980). Eggs are rela¬
tively large, about 1.5 mm in diameter.
The nearly identical Brazilian species, P. nattereri, has been
bred in captivity (Azuma 1975, Brandy & Campbell 1984). Those authors
always found the eggs in the early morning hours, and were usually
guarded by one of the parents. The eggs were pale orange, about 1/16"
in diameter, and had been scattered over an area of 1-1.5 square feet.
Surprisingly for a species with parental care, the eggs were reported
not to be adhesive. The eggs hatched within 48-72 hours and the yolk
sac was absorbed in another 48-72 hours. The relatively large fry were
able to feed immediately on brine shrimp nauplii.
P. caribe is often observed lingering in the shallows, among
aquatic vegetation during the breeding season. The males are probably
territorial at this time. They probably make nests by hollowing out a
sphere amidst the aquatic plants.
Migrations. Apparently this species migrates with the change of
the seasons, downstream at the beginning of the wet season and upstream
at the beginning of the dry season, but little hard data supports this
supposition.
Importance. This species is potentially dangerous to man, but its
"bloodthirstiness" is highly exaggerated. Most reports of "attacks" on
man are probably better explained by scavenging on cadavers of drowning
victims. Piranhas are eaten in some regions, but were not a commercial
species until recently, in response to dwindlinmg stocks of the more
desirable fishes. Many people refuse to eat them because of their sup¬
posed anthropophagous habits.

Pygopristis denticulatus (Cuvier) 1819
Caribe Palometa - Silver Dollar Piranha
525
Figs. 34a (teeth) & 202. Map: fig. 203. Couplet 21a.
Generic synonymy
Pygopristis Müller & Troschel 1844:95 (type species: Pygopristis
fumarius Müller & Troschel 1844 = Serrasalmus denticulatus [Cuvier]
1819, by subsequent designation of Eigenmann 1910); Muller & Troschel
1845:pi. 9, fig. 2a & b.
Specific synonymy
Serrasalmo denticulatus Cuvier 1819:371 (type locality: uncertain);
Günther 1864:367 (British Guiana).
Pygopristis denticulatus Müller & Troschel 1844:95; 1848:637;
Valenciennes in Cuvier & Valenciennes 1849:297; Mago L. 1970:73.
?Pygopristis antoni Fernández Y. 1965:1 (type locality: Río San José,
Guárico state, Venez.); Mago L. 1970:73.
Serrasalmus (Pygopristis! denticulatus Géry 1972:209 (key); 1977:278-80
(key).
Serrasalmus (Pygopristis) antoni Román 1985:111 (Venez.).
Comments. P. antoni Fernández Y. 1965, was described from the Apure
River drainage, Río San José, Guárico state. Géry (1977) stated that the
species described by Fernández Y. (f\ antoni) is more like a Pristobrycon,
but he had not examined any specimens. Fernández Y. described the teeth of
his specimens as follows: "The teeth of this species are tricuspid but
definitely not of the angular type of Serrasalmus and Pygocentrus. They are
of the Pygopri sti s type and their cusps are short, shorter than the width of
their base, which is ellipsoid in cross section. The edges of the cusps are
rounded, not straight, like certain teeth of the Cheirodontids. The third

526
Figure 202
Pyqopristis denticulatus

527
intermaxillary tooth is the shortest, almost typical of the fishes of the
Serrasalmidae group. The mandibular (dentary) teeth are separated at
their bases and do not form a continuous border as in Serrasalmus" (my
translation). He also stated that the first and sixth intermaxillary
(meaning premaxillary ?) teeth are bicuspid in P. antoni, but quadricuspid
and pentacuspid in P. denticulatus. I have compared specimens from the
Aguaro River with some from the Amazonas territory, and found them to be
nearly identical, but I have not seen material from Guyana, the supposed
type locality for P. denticulatus. For now, I consider P. antoni to be a
junior synonym of P. denticulatus. but to be certain Guyanan material must
be compared with the Orinoco specimens.
Etymology. PYGO = "pygmaeus," Latin for small, PRISTIS = saw, from
the Greek; DENTICULATUS = probably refers to the multicuspid teeth.
Description
II lustrations. Figs. 34a (teeth) & 202; Myers 1972:113 ; Géry
1977:280; Román 1985:111 (as Serrasalmus P. antoni).
Di agnosis. This fish looks like a cross between a Metynnis with a
Serrasalmus. The roof of the mouth lacks teeth. The jaws are relatively
weakly developed for a piranha, with incisor-like teeth that are symmetri¬
cal, at least pentacuspid, and not specialized for tearing flesh as in
Serrasalmus species. The anal fin is bilobed in males. Unlike most other
piranhas, which have completely scaled anal fins, this species has at most
two rows of scales on the anal-fin base.
Size. It can grow to about 150 mm SL.
Morphology. The body is discoid.
Counts. DR ii17-19.
Pigmentation. The dorsum is silvery, the ventrum red, and the base
of the caudal fin black. The anal, pelvic and pectoral fins are reddish.

Figure 203. Apure drainage distribution of Pygop
istis denticuiatus.

LA UNIvfMSlDAO O'lf SlCMOF
UNIVERSIDAD NACIONAL E X PE RIME NTAl
DE LOS LLANOS OCCIDENTALES
E ZEQUIE L ZAMORA'
VICE-RECTORADO EN PORTUGUESA
4 .•<
CUENCA DEL RIO APURE
529

530
Distribution and Natural History
Range. It occurs in the Guyana and the Orinoco Basin.
Apure distribution. Map: fig. 203. It is so far known only from
the Aguaro River system.
Habitat. It lives mainly in blackwater streams.
Abundance. RARE.
Number of specimens examined. 4 from 1 collection.
Food. OMNIVORE. It probably feeds on small aquatic animals,
seeds, and fruits etc.
Reproduction. Strategy: r2. It is probably an egg scatterer that
synchronizes reproduction with the onset of the rains in May or June.
Importance. Ornamental.
Roeboides affinis (Giinther) 1868
Dientoncito
Figs. 29a (head) & 204. Map: fig. 205. Couplet 8a.
Generic synonymy
Roeboides Giinther 1864:333 (type species: Ep i cvrtus mi crol epi s
Reinhardt, by subsequent designation of Eigenmann 1910:445); Jordan
1919:333 (designated as logotype Anacyrtus quatemalensis Giinther
[1ater invalidated]).
Rhoeboides Berg 1899:94 (type species: Epicvrtus microlepis Reinhardt).
Cynocharax Fowler 1906 (1907):457 (type species: Anacyrtus affinis
Giinther, by original designation).
Specific synonymy
Anacyrtus affinis Giinther 1868:481 (type locality: Rio Huallaga, Alto
Amazonas), seen; Peters 1877:472 (Calabozo, Venez.).
Cvnopotamus affinis Garman 1890:12.

531
Figure 204. Roeboides affinis.

532
Roeboides affinis Fowler 1905 (1907):457; 1939:269; 1943:100; 1945:163.
Eigenmann 1910:445; Eigenmann & Allen 1942:258 (Orinoco); Mago L.
1970:73.
Roeboides rubrivertex Cope 1872:265 (292) (type locality: between the
mouths of the Rio Negro and the Rio Ucayali).
Comments. Pellegrin (1899) cited Anacvrtus mi crolepis (Rein¬
hardt) 1849 from the Apure River, but that species has over 110 lateral
scales, so the reference is probably a misidentification of one of the
other cynopotamines, perhaps of Cvnopotamus or Charax. Schultz copied
the error (1944b). Román (1985) misidentified Charax qibbosus as
Roeboides mi crolepis.
Etvmology. ROEBOIDES = probably from the Greek "rhomboeides"
which means "resembling a rhombus," in reference to the humped back of
these fishes; AFFINIS = related to, or like.
Description
Illustrations. Figs. 29a (head) & 204; Román 1985:160.
Diagnosis. In the Apure drainage, only Roeboides, Exodon and
Serrabrvcon species have "tusks" or everted teeth, which are used to
scrape scales from other fishes. Exodon paradoxus is easily recognized
by its pigmentation pattern (fig. 32). Serrabrvcon magoi has only 18
anal-fin rays versus over 45 in R. affinis. R. davi has smaller "tusks"
than R. affinis, and differs further in having the humeral spot placed
much farther back on the body (nearer a vertical through the dorsal-fin
origin than the opercle). R. affinis also has more lateral-line scales
than R. davi (69-75 vs 55-61). Small specimens of Roeboides are superfi¬
cially similar to Charax and Cvnopotamus. Upon closer inspection with mag¬
nification, small "tusks" can usually be seen, even in very small Roeboides.

533
Size. It grows to about 70 mm SL.
Morphology. The humpbacked profile of this fish is characteristic.
The upper jaw extends well beyond the lower. The "tusks" are obvious and
well developed.
Counts. AR 47-53; LLS 69-75.
Pigmentation. This fish is nearly transparent in life, but turns
white in preservative. It has a faint humeral spot, and sometimes a faint
caudal spot as wel1.
Distribution and Natural History
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 205. It is common in the lower
11 anos.
Habitat. R. affinis lives in slower-moving lowland streams and in
ponds.
Abundance. COMMON.
Number of specimens examined. 826 from 111 collections.
Food. CARNIVORE. This species employs a highly specialized feeding
adaptation, the "tusks," to scrape scales from the sides of other fishes.
It also includes small aquatic invertebrates and fishes in its diet. It
stalks prey in a head-down position, slowly advancing towards its victim in
a head on position, until it reaches striking distance.
Reproduction. Strategy: rl; fecundity: between 100-750 eggs/ovary;
this egg scatterer breeds all year round, with a peak in reproduction
between February and June (Winemiller & Taphorn 1989).
Importance. This transparent fish is sometimes sold as the "glass
headstander" in the aquarium trade, but its feeding habits quickly make
it unpopular with most aquarists.

Figure 205. Apure drainage distribution of Roeboides affinis.


Roeboides davi (Steindachner) 1878
Dientoncito - Scale-Eating Glassfish
536
Figs. 29b & 206. Map: fig. 207. Couplet 8b.
Specific synonymy
Anacyrtus (Rhaeboides) dayi Steindachner 1878:61 (type locality:
Magdalena Basin, northern Colombia), seen.
Roeboides dayii Eigenmann 1920:12 (Lake Valencia Basin).
Roeboides dayi dayi Schultz 1944b:304; Mago L. 1970: lists.
Roeboides dayi Géry 1977:(key, range).
Comments. Schultz (1944b) described the subspecies R. dayi diento-
nito from the Venezuelan Maracaibo Basin. It supposedly has fewer later¬
al-line scales. However, his sample size was not very large, and the
affinities of the Maracaibo Basin are strongest with the ichthyofauna of
the Magdalena Basin in Colombia, the type locality of R. dayi dayi
Steindachner. Apure drainage specimens might be expected to diverge even
more. It was premature to introduce trinomials into South American fish
nomenclature in 1944b, and still is today in most cases.
Etymology. DAYI = Though I have no direct evidence, this species
might have been named for Benjamin Henry Day (1810-1889) an American jour¬
nalist who published the New York Sun.
Description
Illustrations. Figs. 29b (head) & 206.
Diagnosis. See diagnosis for R. affinis. The upper jaw is about as
long as the lower. The "tusks" are present, but are not as prominent as
in R. affinis, and in small specimens are often hard to see.
Size. It grows to about 60 mm SL.
Counts. AR 50-53 (N=5); LLS 55-61 (N=6).

537
Figure 206. Roeboides davi

Figure 207. Apure drainage distribution of Roeboides dayi.


540
Pigmentation. It is transparent in life with a black humeral spot,
but turns white in preservative.
Distribution and Natural Hi story
Range. It occurs in Venezuela, Trinidad and Colombia.
Apure distribution. Map: fig. 207. This species is very widespread
throughout the piedmont and llanos.
Habitat. This fish is most common in lentic habitats, such as the
slower-moving sections of streams and rivers, as well as in ponds and
swamps, bajios and esteros.
Abundance. ABUNDANT.
Number of specimens examined. 8,067 from 279 collections.
Food. CARNIVORE. It feeds on the scales of other fishes, and
aquatic insects, especially Ephemeroptera (K. Winemiller, pers. com.).
Reproduction. Strategy: rl. This species reaches sexual maturity
quickly (about 3 months). Individuals can breed at least twice a year,
but reproduction occurs in all months. It is probably an egg scatterer.
Importance. Ornamental.
Salminus hi 1arii Valenciennes 1849
Dorado, Saltador - South American Salmon
Fig. 208. Map: fig. 209. Couplet 49a.
Generic synonymy
Salminus Agassiz in Spix 1829:76 (type species: Hvdrocvon brevidens
Valenciennes, by monotypy, (not Hvdrocvon brevidens Cuvier) =
Salminus maxillosus Valenciennes); Müller & Troschel 1845:16.
Specific synonymy
Salminus hi 1 ari i Valenciennes i_n Cuvier & Valenciennes 1849:64 (type
locality: Río Sao Francisco); Pellegrin 1899:157 (Apure River, Venez.)

541
Figure 208. Salminus hi 1 ari i.

542
Schultz 1944b:293 (Aragua state, Venez.) Mago L. 1970:73; Géry
1977:331, 334.
Comments. This species is difficult to distinguish from S. affinis
Steindachner, with which it is supposedly sympatric. Restricted as they
are to montane headwaters, it is hard to understand how a northern South
American species of Salminus could be the same as that in the Río Sao
Francisco of Paraguay and southern Brazil. Despite the popularity of these
fishes as sport and food fish, little has been done on their taxonomy.
Etvmology. SALMINUS = like a salmon. HILARII = meaning unknown.
Description
Illustrations. Fig. 208; Géry 1977:331.
Diagnosis. There are two rows of conical teeth on both jaws, but
none are canines that stand out above the rest. The maxilla is almost
completely toothed. S. hi 1arii might be confused with small Brycon
whitei but the latter differs significantly in dentition, has a much
deeper body, a shorter, more rounded snout, and more lateral-line scales
(66-70 vs 55-65).
Size. It is usually taken at 400-500 mm SL, but are said to reach
nearly a meter.
Morphology. This fish is salmon-shaped with an elongate, fusiform
body that is little compressed. The lateral-line is complete.
Counts. DR 11; AR 24-26; LLS about 60-72; TS 10 above LL.
Pigmentation. Live fish are spectacularly colorful, with bright
orange spots along the sides mixed with blue, yellow and black.
Distribution and Natural History
Range. Though exact boundaries are still uncertain, its range
at least includes the Andean streams of the Orinoco Basin in Colombia

543
and Venezuela. This species has been cited from the Andes as far south
as Bolivia, but those reports are probably based on another species.
Apure distribution. Map: fig. 209. The map reflects more about
fishing efforts with hook and line, rather than the actual distribution of
this species. The Saltador is almost never caught with seines, our most
usual mode of collecting. It is restricted to mountain and piedmont
streams as a rule, though the collection from Bruzual, on the banks of the
Apure River itself, indicates that it can survive in the lowlands, at least
for a while. The specimen in question is a juvenile that probably washed
downstream during the torrential rains typical of the wet season in the
Apure drainage.
Habitat. It is usually seen in montane rivers with strong, fast
currents. It prefers streams with clear, clean water that runs over rocky
substrates. For a fish of this size, it inhabits relatively shallow moun¬
tain river reaches with typical depths of 40 cm.
Abundance. UNCOMMON. The scarcity of specimens reflects more their
ability to avoid capture, rather than actual rarity. We sampled infre¬
quently in the usually inaccessible mountain streams and rivers where this
species lives.
Number of specimens examined. 26 from 15 collections.
Food. CARNIVORE. Piscivore. This species is usually seen in
groups, frequently in mixed shoals with Brvcon whitei, patrolling the
pools just below rapids for potential prey. It will readily take a
silver spoon offered by anglers.
Reproduction. Strategy: r2. It is probably an egg scatterer that
synchronizes reproduction with the onset of the rains in May to June.
Migrations. It is known to migrate annually for great distances.
The Apure populations move upstream in the dry season.

Figure 209. Apure drainage distribution of Salminus hilarii.


546
Importance. This is a magnificent and highly valued sport fish.
Serrabrycon magoi Vari 1986
Sardinita - Scale-Eating Tetra
Fig. 211. Map: fig. 212. Couplet 13b.
Generic synonymy
Serrabrycon Vari 1986:329 (type species: Serrabrycon magoi Vari 1986, by
original designation).
Specific synonymy
Serrabrycon magoi Vari 1986: 329 (type locality: Venezuela, Territorio
Federal Amazonas, Departamento Rio Negro, lower portion of Caño Manu,
which drains into the Rio Casiquiare about 250 m upstream of Solano,
approx. 02° 00’ N, 66° 57’ W), seen.
Comments. This species was recently described from the Amazon Terri¬
tory of Venezuela. Its presence in the blackwater streams of the Apure
drainage point to a link between these two faunas.
Etymology. SERRA = saw, from the Latin, BRYCON = a genus of chara-
cid; MAGOI after Dr. Francisco Mago Léccia, eminent Venezuelan ichthyolo¬
gist.
Description
Illustrations. Fig. 211; Vari 1986:330.
Diagnosis. The overall shape, and anal-fin ray count of 18 easily
separates this species from Roeboides that have over 45 anal -fin rays. The
only other fish in the Apure drainage with "tusks" is Exodon paradoxus, but
it is easily distinguished by the pigmentation pattern (fig. 32).
Size. This is a small tetra, usually seen between 20-40 mm SL.
Morphology. Mammilliform everted teeth or "tusks" are present in
both jaws. The lateral-line is incomplete, with only 7-8 pored scales.

547
Figure 211. Serrabrvcon magoi.

Figure 211. Apure drainage distribution of Serrabrycon magoi.

LA UNIVtASlOAO O'lt Sit MBF
UNIVERSIDAD NACIONAL EXPERIMENTAL
DE LOS LLANOS OCCIDENTALES
"EZEQUIEL ZAMORA"
VICE-RECTOR A DO EN PORTUGUESA
CUENCA DEL RIO APURE
/
MEflIOA A
*
549

550
Counts. DR ii8-ii9; AR iiil5, iv 15 or iv16; PR 12-14; VR i6i; GR
10-11 on the lower limb of the outer arch.
Measurements. GBD 28-30% SL; HL 30-33% SL.
Pigmentation. The caudal-fin base has a distinct black spot. In
live and freshly preserved specimens there is a red dot above and slightly
anterior to the black dot. The base color is tan in preservative. The
scales on the dorsum are edged in gray but have whitish centers. The body
is heavily peppered with melanophores.
Pistribution and Natural History
Range. It occurs in the Orinoco Basin, and is probably present in
the Amazonian Rio Negro drainage as well.
Apure distribution. Map: fig. 212. It is known only from the
extreme southeast corner of the drainage in the Aguaro system.
Habitat. It lives in shady blackwater streams and morichales.
Abundance. RARE.
Number of specimens examined. 7 from 2 collections.
Food. CARNIVORE. Lepidophage. It feeds on the scales of other
fishes, but probably also includes aquatic invertebrates in its diet.
Reproduction. Strategy: rl. It is probably an egg scatterer that
synchronizes reproduction with the onset of the rains in May to June.
Serrasalmus
The genus Serrasalmus is a taxonomic puzzle. Piranhas change shape
and pigmentation as they mature, and have distinctive colors during the
reproductive season. They also seem to be highly variable in coloration
within populations, and among the different tributaries of the Orinoco
Basin. As a result, there are many poorly described species and many
synonyms. The names used here were recommended by Drs. William Fink and

551
Antonio Machado A., who are currently revising the subfamily. Names may
change once they conclude their research. A few of the species reported
here are out of alphabetical order because their names have changed several
times during the preparation of this manuscript. Several nominal species
cited from Venezuelan waters have not been included here; among them we
find: Serrasalmus serrulatus (Valenciennes) 1849; S. spilopleura Kner 1859;
S. scapularis Giinther 1864; S. qvmnoqenvs Giinther 1864; and S. calmoni
Steindachner 1908; S. coccogenis Fowler 1911; Serrasalmus eigenmanni Norman
1928, described from Rockstone, British Guiana; and S. nalseni Fernández Y.
1969 also from Venezuela. Some of these names may eventually replace those
used in this report. Because of these uncertainties very limited synony¬
mies are given for the five Serrasalmus species recognized here.
Serrasalmus altuvei Ramirez 1965
Caribe Azul - Blue Piranha
Fig. 212. Map: fig. 213. Couplet 26a.
Generic synonymy
Serrasalmus Lacepede 1803:283 (type species: Salmo rhombeus Linnaeus,
by monotypy.
Serrasalmo Dumeril 1806:342 (type species: Salmo rhombeus Linnaeus, by
monotypy).
Specific synonymy
Serrasalmus altuvei Ramirez 1965:1 (type locality: Rio San José,
Guárico state, Venezuela), seen; Mago L. 1970:73; Géry 1977:282; Nico
& Taphorn 1986:33; Nico & Taphorn 1988:312.
Types. Holotype: MAC 65.627 (lost?). Paratypes: (MAC 65.627 (7); AFY
65.307 (7); MAC 65.641.

552
Figure 212. Serrasalmus altuvei.

553
Etymology. SERRA = saw, referring to the serrate ventral keel,
SALMUS = salmonlike fish; ALTUVEI = after Dr. Néstor Altuve, the Director
of Natural Resources of the Agriculture Ministry in Venezuela in 1965.
Description
Illustrations. Fig. 212; Ramirez 1965:1; Myers 1972:114; Román
1985:108; Nico & Taphorn 1986:33.
Diagnosis. This piranha can be recognized by the relatively deep
body (its greatest depth measure less than 1.7 times in the SL) and the
very steep dorsal profile. It is most similar to S. rhombeus, but lacks
the massive lower jaw characteristic of that species. It differs further
from that species in having 16-19 (usually 17 or more) vs 15-17, (usually
16 or fewer) dorsal-fin rays.
Size. It grows to at least 170 mm SL.
Morphology. The head is small, and the snout pointed, its dorsal
profile very steep. The eye is about as long as the snout. The body is
quite deep.
Counts. DR Ü14-Ü17; AR 34-38. Ventral scutes 25-31.
Measurements. GBD measuring less than two in SL, (56-67% SL). The
dorsal-fin base is about half as long as the anal-fin base. The adipose
fin is very small, its length contained almost six times in the distance
from the posterior margin of the dorsal-fin base to the posterior margin
of the adipose fin base.
Pigmentation. In preservative the body is heavily marked with irreg¬
ular, vertically elongate dark blotches. The humeral bar is small, trian¬
gular and located just behind the opercle. The thorax and ventrum have no
spots and are lighter colored than the dorsum. The dorsal fin has a faint
dusky blotch. The adipose, caudal and anal fins are margined in black,
with more pigment on the rays, very little on the membrane. The scales in

Figure 213. Apure drainage distribution of Serrasalmus altuvei.

555

556
the humeral area have dark centers. The pectoral and pelvic fins are
transparent (Ramirez 1965).
Distribution and Natural History
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 213. This species just makes it
into the Apure drainage, in the area of the modules, in northern Apure
state. It is probably present in the Aguaro River system as well.
Habitat. It inhabits blackwater streams with abundant aquatic vege¬
tation.
Abundance. UNCOMMON.
Number of specimens examined. 14 from 6 collections.
Food. CARNIVORE. Piscivore. Small juveniles (20-80 mm SL) special -
ize on the fins of other small fishes. As they grow their diet includes
mostly small fish, pieces of fish flesh and fins (Nico & Taphorn 1988).
Reproduction. Strategy: r2. It is probably an egg scatterer that
synchronizes reproduction with the onset of the rains in May to June.
Importance. It is potentially valuable as an ornamental.
Serrasalmus medinai Ramirez 1965
Caribe - Piranha
Fig. 214. Map: fig. 214. Couplet 28b.
Specific synonymy
Serrasalmus medinai Ramirez 1965:1 (type locality: El Polvero, Rio San
José, estado Guárico, Venezuela).
Pristobrvcon sp Nico & Taphorn 1986:33 (diet).
Serrasalmus caribe Nico & Taphorn 1988:311 (diet).
Types. Holotype: MAC 65.622 (lost?). Paratypes: MAC 63.604, AFY
63.200 (lost?).

557
Figure 214. Serrasalmus medinai

558
Comments. This species is presented out of alphabetical order be¬
cause it was recently reidentified by Drs. W. Fink and Machado Allison
(pers. com.), after the species had been listed and the figures and maps
already drawn.
Etymology. MEDINAI = after Dr. Gonzalo Medina, chief of the Division
of Fauna, Natural Resources, Ministry of Agriculture and Animal Husbandry
in 1965.
Description
Illustrations. Fig. 214; Myers 1972:119; Nico & Taphorn 1986:33 (as
Pristobrvcon sp.).
Di agnosis. This piranha is easily confused with S. rhombeus, or S.
altuvei, and specimens must be examined with care to make a correct identi¬
fication. Ramirez (1965) distinguished them as follows: S. al tuvei has
34-36 anal-fin rays, vs 30-31 in S. medinai. Maximum body depth is 49-56%
SL in S. medinai. vs 56-67 % SI in S. altuvei. In both S. rhombeus and S.
altuvei the adipose fin is short, its length contained more than three
times in the space between the dorsal fin and the adipose; in S. medinai it
is longer, barely fitting twice. Another good character to distinguish them
is the shape of the teeth on the roof of the mouth which are unicuspid and
squared off distally, but with a more-or-less conical base and usually
number fewer than five. The caudal fin is almost always edged in black (as
in S. rhombeus). There is no black line along the base of the anal fin,
but this and the dorsal fin are often edged with black. The sides are
heavily spotted, and there is a dark humeral spot behind the opercle.
Size. It reaches at least 200 mm SL.
Morpholoqv. This species is intermediate in build, between the
massive Pygocentrus caribe and the more slender Serrasalmus rhombeus.

559
Counts. DR 17-20; AR 27-31. Ventral scutes 20-21 to pelvic fins,
31-32 total.
Measurements. (from Ramirez 1965). Eye relatively small, its
diameter fitting about 1.5 times in the snout length. Dorsal-fin base
length contained less than 1.7 times in anal-fin base. GBD 49-56% SL. HW
18-28% HL.
Pigmentation. See diagnosis. The humeral spot is distinct, triangu¬
lar, and located just behind opercle. The caudal fin is almost always
edged with black (less frequently the dorsal anal, and adipose fins are
edged in black as well). The body is silvery in life, but specimens pre¬
served in formalin often have diffuse circular spots arranged in horizontal
lines (Ramirez 1965).
Pistribution and Natural Hi story
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 215. This species is found in the low
llanos.
Habitat. It lives in lowland llanos streams and rivers.
Abundance. COMMON.
Number of specimens examined. 216 from 68 collections.
Food. CARNIVORE. Piscivore. The adults bite fins from large fishes
and eat smaller whole fish as well. Subadults eat fins and aquatic in¬
sects, but smallest juveniles feed on microcrustaceans (Nico & Taphorn
1986, 1988; K. Winemiller pers. com.).
Reproduction. Strategy: r2; fecundity: 3048 eggs/female; egg diame¬
ter 1.5 mm (Winemiller & Taphorn 1989). It is probably an egg scatterer.
Sexual maturity is reached in one year. Spawning occurs in the first three
months of the rainy season. Individuals may spawn more than once.

R g tire 215. Apure drainage distribution of Serrasalrrus medinai.


562
Importance. This fish is only eaten when other more desirable spe¬
cies aren’t available. In general, piranhas are not eaten in the llanos
because, according to popular belief, they sometimes include humans in
their diet.
Serrasalmus elongatus Kner 1860
Caribe Alargado, Caribe Pinche - Elongate Piranha
Fig. 216. Map: fig. 217. Couplets 25a & 27a.
Specific synonymy
Serrasalmus elongatus Kner 1860:44, pi. 5, fig. 12 (type locality: Rio
Guaporé, Mato Grosso); Géry 1977:282-3, 285; Nico & Taphorn 1986:24;
Nico & Taphorn 1988:311.
Serrasalmus pingke Fernández Y. 1951:1 (type locality: Río Apure, La
Defensa, al S.O de San Fernando, Venezuela).
Comments. Géry (1977) stated that Serrasalmus pingke Fernández Y.
1951 could be a synonym of this species. Since S. pingke was described
from two juveniles specimens of only 55.4 and 58 mm SL, it is not possible
to determine if it is identical with the Amazonian S. elongatus. Dr.
Antonio Machado (pers. com.) is currently investigating the possibility
that there are two elongate piranhas in the Orinoco Basin. If this should
be the case, S. pingke might prove to be a valid species.
Etymology. ELONGATUS = elongate.
Description
Illustrations. Fig. 216; Norman 1928:806; Fernández Y. 1951:1 (as
S. pingke); Géry 1977:282; Román 1985:106, (as S. pingke); Nico & Ta¬
phorn 1986:33.
Diagnosis. The relatively elongate body (for a piranha) is suffi¬
cient to identify this species.

563
Figure 216.
Serrasalmus elonqatus.

564
Size. It reaches at least 250 mm SL.
Morphology. This is the most elongate piranha in Venezuela.
Counts. DR ii13.
Measurements. GBD 42% SL.
Pigmentation. The opercle has a dark smudge at the angle. The
humeral spot is triangular with the point down, and is located behind the
opercle. The adipose fin is pigmented at its tip. The caudal fin is dark.
In life this species is plain silvery most of the year. In the breeding
season, the chest becomes reddish. When it is preserved the dorsum and
upper sides have numerous darker flecks and blotches that often run togeth¬
er to form vertical bars, especially in the humeral area.
Pistribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map: fig. 217. It is known only from northern
Apure state and the Aguaro River system.
Habitat. It lives in blackwater streams of the low llanos.
Abundance. UNCOMMON. Adults of this species are seldom collected
with seines, so their apparent scarcity could be an artifact of our col¬
lecting methods. Most large specimens were captured on hook and line.
Number of specimens examined. 51 from 17 collections.
Food. CARNIVORE. Piscivore. The adults clip fins from large
fishes and also eat whole small fish, subadults eat fins and aquatic
insects, smallest juveniles take microcrustaceans (Nico & Taphorn 1988;
K. Winemiller pers. com.).
Reproduction. Strategy: r2. It is probably an egg scatterer that
synchronizes reproduction with the onset of the rains in May to June.
Importance. It is consumed locally.

Figure 217. Apure drainage distribution of Serrasalmus elongatus.


567
Serrasalmus irritans Peters 1877
Caribe Pinche - Slender Piranha
Fig. 218. Map: fig. 219. Couplet 27b.
Specific synonymy
Serrasalmo irritans Peters 1877:472 (type locality: San Fernando de
Apure, Venezuela).
Serrasalmus irritans Eigenmann & Eigenmann 1891:60; Schultz 1944b:255
(as synonym of S. spilopleura); Nico & Taphorn 1988:312.
Serrasalmus eigenmanni: Schultz 1944b:255 (Rio Caripe, Venez.); Taphorn
& Lilyestrom 1983:70 (Apure modules, Venez.); Nico & Taphorn 1986:31,
33 (Apure drainage, Venez.).
Comments. Doubt still exists as to the true identity of this species
(as for most piranhas), but at least it was described from the Apure drain¬
age. I have called this species S. eigenmanni in previous works. There
may be a complex of species involved here.
Etymology. IRRITANS = bothersome.
Description
Illustrations. Fig. 218; Román 1985:108; Nico & Taphorn 1986:31,33
(as Serrasalmus eigenmanni).
Diagnosis. This species is extremely compressed. It has a very
slender head (the least interorbital width measures more than 3.3 times
in the head length). The black markings along the base of the caudal
fin (fig. 218) are also characteristic of this species. It is further
distinguished by having 32 or more anal-fin rays.
Size. It grows to about 150 mm SL.
Counts. DR i i 13; AR 34.
Pigmentation. In preservative these fishes have the upper sides
flecked with numerous black spots. The caudal fin is black along its
base and the anal fin is sometimes darkly pigmented as well.

568
Figure 218. Serrasalmus irritan*.

Figure 219. Apure drainage distribution of Serrasalmus irritans.


571
Distribution and Natural History
Range. It is known only from the Orinoco Basin.
Apure distribution. Map: fig. 219. It is common in the lower lla¬
nos, but is also found in the piedmont.
Habitat. It inhabits lowland llaneran streams.
Abundance. COMMON. This is usually the second most abundant piran¬
ha in most biotopes (after Pygocentrus caribe). It is widespread through¬
out the drainage.
Number of specimens examined. 744 from 142 collections.
Food. CARNIVORE. Piscivore. Adults mostly take whole small fish
but clip fins from large fishes as well. Juveniles specialize in eating
fins, which are frequently the only item in their stomachs. Small aquatic
insects and microcrustaceans comprise the diet of the smallest juveniles
(Nico & Taphorn 1986, 1988; K. Winemiller pers. com.).
Reproduction. Strategy: r2; fecundity: 280 eggs/female; egg diame¬
ter: 1 mm. Sexual maturity is reached in one about year. Spawning occurs
during the first three months of the rainy season. It is probably an egg
scatterer (Winemiller & Taphorn 1989).
Importance. It is consumed locally.
Serrasalmus rhombeus (Linnaeus) 1766
Caribe Blanco, Caribe Mondonguerro, Caribe Ojo Rojo
White Piranha, Red-Eye Piranha
Fig. 220. Map: fig. 221. Couplet 26b.
Specific synonymy
Salmo rhombeus Linnaeus 1766:514 (type locality: Surinam), seen.
Serrasalmus rhombeus Mago L. 1970:73; Géry 1977:283, 286, 288-91; Novoa
1982:273 (biology); Machado A. & Garcia 1986:193 (diet); Nico &
Taphorn 1986:24; Nico & Taphorn 1988:312.

572
Figure 220. Serrasalmus rhombeus.

573
Comments. Géry (1977) stated that S. medinai Ramirez 1965 is proba¬
bly a synonym of this species. The name S. rhombeus is used for forms that
range from Surinam and the Guianas, as well as the Orinoco and Amazon
basins. Further research will probably justify taxonomic recognition of
some of these geographically isolated populations. In the Orinoco drainage
it is possible that a complex of species is included under this name.
Etymology. RHOMBEUS = probably refers to their rhomboid shape when
older.
Description
Illustrations. Fig. 220; Myers 1972:122; Géry 1977:288, 291;
Novoa et al. 1982:fig. 9; Román 1985:110; Axelrod et al. 1971:F-556.05,
556.06, 559.00; Nico & Taphorn 1986:40.
Diagnosis. In contrast to the situation observed in S. caribe with
which this species is most often confused (see diagnosis of that species),
the teeth on the roof of mouth, are triangular, bi- to tricuspid (the
lateral cusps are small and hard to see), and usually number seven or more.
The caudal fin is usually edged with black. There are 15-17 (usually 16 or
17) dorsal-fin rays. The spots on the sides are usually rounded, and not
vertically elongated. Specimens over 150 mm have a massive lower jaw; the
jaw width at the commissure is greater than the eye diameter.
Size. Novoa et al. (1982) reported a specimen of 295 mm TL that
weighed 0.47 kg. Axelrod et al. (1971) claimed that this species can reach
420 mm TL.
Morphology. This species changes shape considerably as it grows.
Young specimens are compressed and relatively elongate. As they grow they
become stockier, and have a relatively deeper body. Géry (1972) discussed
these allometric changes at length.

574
Counts. DR i i 13; AR 33.
Pigmentation. This species changes its pigmentation as it grows.
Young specimens are basically whitish or yellowish with a silver-gray
background color on the body, and a reddish abdomen. The fins are hyaline
except for the caudal fin which is edged in black, and the pectoral fins
which are reddish. Individuals darken progressively until they turn com¬
pletely black with purple and gold flecks on the flanks. The fins darken
along with the rest of the body, and eventually can become all black. The
eye is usually bright red at all sizes.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins of Brazil,
Venezuela and Colombia, and in Guyana and Surinam.
Apure distribution. Map: fig. 221. Curiously, this species has
not been taken in the far western portion of the drainage. It is found
throughout the lower portions of the rest of the drainage.
Habitat. S. rhombeus lives in lowland streams throughout the
drainage, but is more common in black and Clearwater streams.
Abundance. COMMON.
Number of specimens examined. 136 from 61 collections.
Food. CARNIVORE. Piscivore. This species is known to feed both
during the day and at night. Adults bite fins from large fishes and eat
small whole fish, subadults eat fins and aquatic insects, and the
smallest juveniles eat microcrustaceans (Nico & Taphorn 1988; K.
Winemiller pers. com.).
Reproduction. Strategy: r2; fecundity: 4303 eggs/female; egg diame¬
ter: 2.25 mm (Winemiller & Taphorn 1989). Novoa et al. (1982) found matur¬
ing females in the Orinoco Delta in March. In the llanos spawning occurs

Figure 221. Apure drainage distribution of Serrasalmus rhombeus.

en
\i
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577
during the first three months of the rainy season. Sexual maturity is
reached in one year. Individuals may spawn more than once.
Importance. It is consumed locally.
Tetragonopterus argenteus Cuvier 1817
Sardina - Silver-Foil Tetra
Fig. 54 & 222. Map: fig. 223. Couplet 84a.
Generic synonymy
Tetragonopterus Cuvier 1817:166 (type species: Tetragonopterus argenteus
Cuvier, by monotypy).
Specific synonymy
Tetragonopterus argenteus Cuvier 1817 (1818):455 (type locality: Bahia,
Brazil); Pellegrin 1899:157; Schultz 1944b:338; Mago L. 1970:73; Géry
1977:448-50, 454.
Etymology. TETRA = four, GONO = angle, PTERUS = wing or fin, thus
meaning square winged; ARGENTEUS = silvery.
Description
Illustrations. Fig. 54 & 222; Eigenmann 1917:pi. 2, 4; Géry
1977:448, 449; Román 1985:177; Axelrod et al. 1971:F-578.21, 578.22.
Diagnosis. The decurved lateral-1ine, deep body, and relatively
large eye are characteristic of this species. The other species in the
genus T. chalceus is present in neighboring drainages, and was reported by
Pellegrin (1899) from the Apure River. It is less deep bodied, has only 3-
4 scales bordering the occipital process (vs 5-7), and has fewer anal rays
32-33 (vs 36-37 in T. argenteus).
Size. It grows to about 110 mm TL, but most individuals are smaller,
between 40-70 mm SL.

578
Figure 222. Tetraqonopterus arqenteus

579
Morphology. The body is deep and greatly compressed, with a steep
upward curve to the dorsal profile. The complete lateral line is de-
curved, and passes close to the anal-fin base.
Counts. DR 11; AR 36-37; LLS 31-35; TS 9 above LL, 4-5 below.
Measurements. The eye is very large, measuring 3.0 times in the head
length in the adult, and 2.25 in young.
Pigmentation. The body is silvery with a double humeral bar, and a
large caudal blotch. The pelvic fins and anterior anal fin are red.
Distribution and Natural History
Range. It occurs in the Amazon, Orinoco and La Plata basins, as
well as in Guyana.
Apure distribution. Map: fig. 223. It is found throughout the
11 anos.
Habitat. This fish is usually found in slower parts of streams and
rivers.
Abundance. COMMON.
Number of specimens examined. 178 from 35 collections.
Food. OMNIVORE. It feeds on seeds, aquatic insects, and fish car¬
rion (K. Winemiller, pers. com.).
Reproduction. Strategy: r2; fecundity is high with 3398 eggs/ fe¬
males; egg diameter: 1.00 mm (Winemiller & Taphorn 1989). Sexual maturity
is reached in about one year. Spawning occurs during the first month of
the rainy season. Individuals usually spawn only once per year (Winemill¬
er & Taphorn 1989). Sexual dimorphism is evident. Males are smaller and
slimmer and with prolonged dorsal-fin rays in older, more mature specimens
(Axelrod et al. 1971).
Importance. Ornamental.

Figure 223. Apure drainage distribution of Tetragonopterus argenteus.

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582
Triportheus
I have identified what I believe to be three species of Triportheus
among the collections made in the Apure drainage for this report, but
assigning correct names to the three has proven difficult. The differences
are presented in the key, but there is so much variation in color and
meristics that I suspect the situation may be more complex. Perhaps hybri¬
dization is involved. At any rate, the three forms are: a red-tailed, very
deep-bodied species, called here T. sp. "cola roja" (meaning red tail in
Spanish) and I. rotundatus by other authors; a species that usually has a
black stripe though the tail, but can also have some red in the tail,
I. anquíatus: and an elongate, yellow-tailed form T. albus found so
far only in main river channels. All seem to grade into one another.
Triportheus albus Cope 1872
Arenca Cola-amarilla - Yellow-Tailed Triportheus
Fig. 224. Map: fig. 225. Couplet 75b.
Generic synonymy
Triportheus Cope 1872:264 (type species: Triportheus flavus Cope, by
subsequent designation of Jordan 1919:363); Myers 1940:170 (suppression
of Chalcinus).
Chaicinus (non Rafinesque 1815) Valenciennes in Cuvier & Valenciennes
1849:258 (type species: Chalcinus brachipomus Valenciennes, by
subsequent designation of Eigenmann 1910:440).
Specific synonymy
Triportheus albus Cope 1871:264, pi. 14, fig. 3 (type locality: Ambyiacu
River, eastern Ecuador), seen; Géry 1977:336, 343.
Comments. Several authors have reported I. elongatus from Venezuela
(Steindachner 1879; Eigenmann & Eigenmann 1891; Fowler 1911; Eigenmann &

583
Figure 224. Triportheus albus.

584
Allen 1942; Schultz 1944b), but none of these records are specifically
from the Apure drainage. Perhaps their identifications were correct
but it is also possible that these refer to this species. The genus is
in need of complete revision.
Etymology. TRI = three, PORTHEUS = "openings" or "holes" from the
Latin "portus" meaning openings or holes; ALBUS = white, Latin.
Description
Illustrations. Fig. 224; Géry 1977:336.
Diagnosis. The caudal fin is usually yellow. The body is elongate
and slender, its greatest depth measures 3.4 times in the SL. There are
usually only five (sometimes six) scales between the lateral line and
the dorsal-fin origin.
Size. It can reach 120 mm SL, but most individuals are smaller.
Morphology. The body is elongate, and highly compressed with a
keeled breast. The pectoral fins are placed low on the body and are ex¬
panded and stiff.
Counts. There is one tooth behind the main row on the dentary.
Pigmentation. The body is silvery and the tail yellow.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map: fig. 225. It has been taken only from
big rivers.
Habitat. T. albus seems to be restricted to big rivers.
Abundance. RARE.
Number of specimens examined. 181 from 5 collections.
Food. OMNIVORE. It feeds on seeds, terrestrial insects, and
aquatic insects. Goulding (1980) noted that the species that live in

Figure 225. Apure drainage distribution of Triportheus albus.

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587
larger rivers and feed on potamoplankton have finer gill rakers than the
other Amazonian species. This is not the case for this species, which has
gillrakers of about the same size as the other two.
Reproduction. Strategy: r2; fecundity: about 3000 eggs/female; egg
diameter: 1.8 mm (Winemiller & Taphorn 1989). It is probably an egg scat-
terer. Spawning occurs annually during the first month of the rains.
Importance. It is often used as bait by commercial fishermen.
Triportheus anquíatus (Spix) 1829
Arenca - Triportheus
Fig. 226. Map: fig. 227. Couplet 75a.
Specific synonymy
Chalceus angulatus Spix j_n Agassiz 1829:67, pi. 34 (type locality: "in
Brasiliae aequinoctialis fluviis"), seen.
Chalciñus angulatus Steindachner 1879:157 (Ciudad Bolivar); Eigenmann &
Eigenmann 1891:56 (Orinoco); Eigenmann & Allen 1942:261 (Orinoco
Basin); Schultz 1944b:274; Mago L. 1970:73; Géry 1977:332-3, 343, 346.
Etvmology. ANGULATUS = probably referring to the sharp angle of the
sharply keeled chest.
Description
Illustrations. Fig. 226; Weitzman 1960:fig. 1; Géry 1977:333.
Román 1985:154.
Diagnosis. The caudal fin is usually red, sometimes with the central
rays black. The body is deeper in this species than in the other Apure
drainage Triportheus spp., its depth measures less than three times in the
SL. There are usually six scales between the lateral line and the dorsal-
fin origin.
Size. It can reach some 200 mm SL, but is usually found smaller.

588
Figure 226. Triportheus angulatus.

589
Morphology. The body is elongate, and highly compressed with a
keeled breast. The pectoral fins are set low on the body, and are enlarged
and stiff. The swim bladder is two chambered, with the anterior chamber
rounded, the posterior one longer and tapering to a point.
Counts. DR 11-12; PR 12; AR 31-36; LLS 32-34; TS 6 (above lateral
line). The dentary has one large tooth behind the main row.
Measurements. GBD 35-43% SL.
Pigmentation. This species is extremely variable. Usually the base
color is silvery, but the upper half of the fish is sometimes dark brown
and the lower half whitish. There is a dark black lateral stripe in some
populations that continues uninterrupted onto the extended central caudal
rays found in some individuals.
Distribution and Natural History
Range. It occurs in the Amazon and Orinoco basins.
Apure distribution. Map; fig. 227. It is known from throughout
the lower llanos.
Habitat. It prefers flowing waters and is found in most llaneran
streams.
Abundance. COMMON.
Number of specimens examined. 105 from 39 collections.
Food. OMNIVORE. It feeds on seeds, terrestrial and aquatic
insects, and small fishes. One 135 mm SL specimen from the Rio Tucupi-
do contained a 13.7 mm SL Creagrutus in its stomach. Goulding (1980)
noted that in Brazil,the deeper-bodied species I. angulatus, included
more terrestrial insects in its diet than the sympatric, more elongate
species I. elongatus. He supposed that the deeper body allowed I.
angulatus to dash more quickly up to the surface to catch fallen prey.

Figure 227. Apure drainage distribution of Triportheus angulatus.

CUENCA DEL RIO APURE
(«CALA AAASCl
too
en
vo

592
Reproduction. Strategy: r2; fecundity: about 3000 eggs/female; egg
diameter: 1.8 mm. It is probably an egg scatterer. Spawning occurs annu¬
ally during the first month of the rains (Winemiller & Taphorn 1989).
Migrations. During the dry season it is often observed below the
tailraces of dams along with Prochilodus mariae, attempting to migrate up¬
stream. It normally would descend again at the end of the dry season in
rivers that haven’t been impounded.
Importance. It is used as bait by commercial fishermen.
Triportheus sp. "cola roja"
Arenca Cola Roja - Red-Tailed Triportheus
Fig. 228. Map: fig. 229. Couplet 74a.
Specific synonymy
?Triportheus brachypomus Pellegrin 1899:157 (Apure River); ?Fowler
1931:408 (Caño Guanoco, Venez.).
?Triportheus rotundatus Schultz 1944b:274.
Comments. This could be T. rotundatus (Schomburgk) 1841 from British
Guiana, but the coloration, counts and measurements given don’t exactly
agree with that species.
Description
Illustrations. Fig. 228; Román 1985:155.
Diagnosis. The caudal fin is usually red in the species, but is
quite variable and often has the central rays black. There are usually
seven scales between the lateral-line and the dorsal-fin origin.
Size. It can reach 200 mm SL, but most individuals are smaller.
Morphology. The body is elongate and highly compressed. The
breast is keeled. The enlarged, stiff pectoral fins are placed low on
the body.

593
Figure 228
Triportheus sp. "cola roja

Figure 229. Apure drainage distribution of Triportheus sp. "cola roja

CUENCA DEL RIO APURE
■ «CALA «AAfCA
en
en

596
Counts. LLS 34-36; TS 7-8 (above lateral line); AR 31-37. The
dentary has one large tooth behind the main row.
Measurements. GBD 34-40% SL.
Pigmentation. The body is silvery and the tail is red, sometimes
with the central caudal rays dark.
Distribution and Natural History
Range. It occurs in the Orinoco Basin, and perhaps Guyana.
Apure distribution. Map: fig. 229. It seems to be absent from the
western portion of the drainage, but are otherwise common in the lower
parts of the drainage.
Habitat. It lives in flowing waters throughout the eastern portion
of the drainage.
Abundance. COMMON.
Number of specimens examined. 1,045 from 107 collections.
Food. OMNIVORE. It feeds on seeds, and on terrestrial and aquatic
insects.
Reproduction. Strategy: r2; fecundity: about 3000 eggs/female; egg
diameter: 1.8 mm (Winemiller & Taphorn 1989). It is probably an egg scat-
terer. Spawning occurs annually during the first month of the rainy sea¬
son.
Migrations. See the previous account.
Importance. It is often used as bait by commercial fishermen.
Xenagoniates bondi Myers 1942
Sardinita Transparente - X-ray Tetra
Fig. 230. Map: fig. 231. Couplet 3a.
Generic synonymy
Xenagoniates Myers 1942:90 (type species: Xenagoniates bondi, by
monotypy).

597
Figure 230. Xenagoniates bondi.

598
Specific synonymy
Xenaqoniates bondi Myers 1942:90, fig. 1 (type locality: Río Amana, 6 km
E of Santa Barbara and 35 km W of Maturin, Monagas state, Venezuela),
seen; Schultz 1944b:312; Mago L. 1970:73; Géry 1977:341, 347.
Types. Holotype: CAS-SU 36486. Paratopotypes: CAS-SU 46487 (3).
Etymology. XENA = from the Greek for strange, GONIATES = probably
from the Greek "gonia" meaning angled, perhaps in allusion to the unusual
shape of this fish, there is also a genus of characid Agoniates which may
have been used as a root; BONDI = after F. Bond, who worked on a mosquito
control program for in Venezuela and collected the holotype.
Description
Illustrations. Fig. 230; Géry 1977:341.
Diagnosis. The posterior position of the dorsal fin and the extreme¬
ly long anal fin (with more than 55 rays) are characteristic of this spe¬
cies. An adipose fin is present.
Size. This is a small fish that seldom exceeds 60 mm SL.
Morphology. This monotypic genus is characterized by having teeth on
the ectopterygoid bone, a very elongate body and a complete lateral line.
The anal fin is long and broad, the dorsal fin is set very far back on the
body. The base of the anal fin has a sheath of scales.
Counts. DR ii8; AR 61-68; PR i12; VR i5; LLS 49-51; TS 13; maxilla,
premaxilla and dentary each with a single row of small tricuspid teeth.
Pigmentation. It is mostly transparent in life, but turns pale
white or yellow in preservative. The occiput, first rays of the
pectoral fin, the basal part of the first dorsal rays, and a median
stripe through the center of the caudal peduncle and caudal fin, are all
darkened with crowded melanophores (Myers 1942).

Figure 231. Apure drainage distribution of Xenagoniates bondi.

10* -
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o
o

601
Distribution and Natural History
Range. It occurs in Venezuela and Colombia, probably throughout tito
Orinoco Basin.
Apure distribution. Map: fig. 231. It is found throughout the
llanos and lower Andean piedmont.
Habitat. It inhabits the slower moving sections of streams and
rivers of the low and high llanos, up to the piedmont region, and occurs
in both white and blackwater biotopes.
Abundance. COMMON.
Number of specimens examined. 574 from 87 collections.
Food. These fishes nip pieces of fins from other fishes.
Reproduction. Strategy: rl. It has a very long extended ovary which
extends back to the end of the anal-fin base, below and between the tips of
the ribs. Eggs are visible through the transparent body wall.
Importance. It is sometimes sold as an ornamental, but its fin
nipping habits make it unpopular.
Unidentified
Fig. 232. Map: fig 233. Couplet lib.
Comments. This fish superficially resembles species of the glandule
caudine genus Prionobrama. which has not yet been recorded from the Orinoco
Basin. Prionobrama, however, has only one row of premaxillary teeth, vs
two in this fish. I suspect it may be a Hemiqrarnrnus, but has the dorsal
fin farther back than most species of that genus.
Description
Illustrations. Fig. 232.
Diagnosis. This fisr '-ese~s'e: Gepr /rocrarax /a e.'o ae out tr.at
species has the dorsal fin placed fa'' posterior to the anal-fin origin.

602
Figure 232. Unidentified.

603
In this unidentified species, the anal-fin origin is almost directly
under the dorsal-fin origin. In Hemigrammus species, the dorsal-fin
origin is anterior to the anal-fin origin.
Size. It reaches about 25 mm SL.
Morphology. The mouth is oblique, and nearly superior, with the
maxilla in a nearly vertical position. The dorsal profile is straight,
as in Gephvrocharax valenciae. The lateral line is incomplete. The
pectoral fins are set quite low.
Counts. DR 9; AR 25-26; PR 10; VR 7; LS 32-34.
Pigmentation. There is a line of melanophores along the dorsal
midline. The area above the anal fin also has scattered dark dots. Other¬
wise it is plain. In live specimens, the body is silvery to bluish and the
tail is pink to red.
Distribution and Natural Hi story
Range. It has been taken only from the upper Rio Orituco and the
neighboring Río Uñare drainage.
Apure distribution. Map: fig. 233. It is known only from the north-
easternmost corner of the Apure drainage, in the Orituco system.
Habitat. The Rio Orituco is a muddy-bottomed river, that is
reduced to isolated pools in the dry season.
Abundance. RARE. This species is abundant where it occurs.
Number of specimens examined. 219 from one site.
Food. OMNIVORE. It probably feeds on aquatic insects, small seeds,
microcrustaceans and other small invertebrates.
Reproduction. Strategy: rl. It is probably and egg scatterer that
synchronizes reproduction with the onset of the rainy season in May or
June, as is true of most small characids.

Figure 233. Apure drainage distribution of an unidentified characid.

10"
CUENCA DEL RIO APURE
ttCALA lliftt
100
605

606
Characidiidae
Members of this family are ecological equivalents of the darters
of North America. They are all small, benthic micro-carnivores, well
adapted to life in the current, among rocks, gravel or sand. The pec¬
toral fins and often the pelvics are enlarged, with the anteriormost
rays thickened for added strength, and the chest flattened to give
maximum contact with the substrate, which allows these fish to appress
themselves closely in order to avoid the rush of the current. The snout
and head are narrow and pointed. The entire body is hydrodynamically
designed for little resistance to flow. Some species bury themselves in
the sand.
Species of Characidiidae are remarkably uniform in morphology, and
many species differ only slightly in outward appearance. As a result,
for many years almost all Characidium were automatically referred to the
species C. fasciatum. The work of Travassos in Brazil (eg. 1967) has
revealed much more diversity. Géry (1977) estimated about "50 or more"
species for the family.
The correct taxonomic rank of the Characidiidae is still unclear.
Schultz (1944b) included them along with most other characoids in the
family Characinidae. Weitzman (1962) still recognized only one family,
the Characidae. The group was split into 16 families by Greenwood et
al_. (1966) but no specific mention of the genus Characidium and its
relatives was made. Mago L. (1970) included seven Venezuelan species
in the family Lebiasinidae, but Weitzman and Cobb (1975) did not include
Characidium and relatives in that family. Weitzman & Kanazawa (1976)
stated that the subfamily Characidiinae is of uncertain family affini¬
ties, tentatively assigned it to the Characidae, and rejected Géry’s

607
(1971) recognition of the Characidiidae (with the subfamilies Chara-
cidiinae, Elachocharacinae and Geisleriinae). Weitzman & Géry (1981)
"tentatively" recognized Characidiinae as a characid subfamily, but
rejected Geisleriinae and Elachocharacinae as junior synonyms. Weitzman
(1986) continues to describe new species as members of the family Chara-
cidae. I have recognized the group at the family level because I believe
the morphological and ecological characters that unite them as a group
will prove sufficient to separate them from the other families of Chara-
ciformes.
Mr. Paulo Buckup, a Brazilian ichthyologist working with Dr. Wil¬
liam Fink at the University of Michigan, is currently revising the
family and will undoubtedly increase the known number of species. He
has generously provided me with his preliminary key to the Venezuelan
species and has also verified or corrected many of my identifications.
I have adopted his species identifications and codes. The species are
out of alphabetical order in the species account because I have pre¬
served the original order based on different (my) code names and species
identifications.
Though little is known about characidiid reproductive biology,
they probably are r2. Work done on the most common species in the
piedmont suggests that they attain sexual maturity in about 11 months,
and that reproduction is repeated throughout the first 4-5 months of the
wet season and longer if conditions are favorable (Winemiller & Taphorn
1989). Hoedeman (1975) reported that "Characidium fasciatus scattered
its eggs amongst aquatic plants near the bottom in an aquarium. Some
North American darters display spectacular nuptial colors, and deposit
small numbers of eggs under rocks, which are then guarded. South Ameri¬
can darters might be expected to show similar adaptations.

608
Key to the Species of Characidiidae in the Apure River Drainage.
la. Dorsal fin with fewer than twelve rays... ...2
lb. Dorsal fin with more than 16 rays...Elachocharax (2 spp.) ...11
2a. (la) Mouth ventral (fig. 234a); body very elongate, its maximum
depth about 12% SL; maxilla with up to ten teeth; snout very elon¬
gate and narrow... Ammocryptocharax elegans (fig. 235)
2b. Mouth terminal or subterminal (fig. 234b & c); body deeper, its
maximum depth greater than 17% SL; no maxillary teeth; snout
deeper and less pointed... ...3
3a. (2b) Caudal fin with distinct vertical bars, sometimes formed by
rows of spots; first four pectoral-fin rays unbranched; body very
dark (black or dark brown), with numerous spots and blotches...
Characidium sp. D (fig. 239)
3b. Caudal-fin not vertically barred, sometimes with a basal spot or
central rays darker or with spots not forming distinct bars; first
three (or fewer) pectoral-fin rays unbranched; body pigmentation
variable, but never dark brown or black... ...4
4a. (3b) Area between the pectoral-fin bases naked, abdomen scaled...
Characidium voladorita (fig. 251)
4b. Both the area between the pectoral-fin bases and the abdomen
scaled... ...5
5a. (4b) Sides without well-defined black vertical bars, a horizontal
row of dark dots or small irregularly arranged crescent-shaped
marks down midside often present, but these never forming distinct
vertical bars along sides; dorsum crossed by narrow brown saddles
that do not continue on sides; lateral line incomplete 6

609
Figure 234. Mouth position in Characidiidae.
a. Ventral mouth position.
b. SubterminaT mouth position.
c. Terminal mouth position.

610
5b. Sides with distinct vertical bars; a dark lateral stripe sometimes
present; dorsum without brown saddles; lateral line variable 7
(Note: In large adults of some species the vertical bars may be
obscured by dark pigmentation, at least during the reproductive
season, but smaller individuals should show this characteristic
at all times.)
6a. (5a) Middle of sides with horizontal row of black spots; gape even
with or above a horizontal line from along ventral edge of eye
(fig. lc); body heavily pigmented with tiny chromatophores that
give a dusky background color in preservative; usually seven
pectoral rays; greatest body depth 20% SL; head length 17% SL;
interorbital width 40% HI Characidium sp. K (fig. 243)
6b. Sides with irregularly arranged small brown crescents; gape below
a horizontal line from along ventral edge of eye (fig. lb); body
nearly immaculate except for light brown crescents on sides; 8-10
pectoral rays; body more elongate, its greatest depth about 16%
SL; head length 25% SL, interorbital 30% HL...
Characidium sp. J (fig. 245)
7a. (5b) Pectoral rays 12-15 ... Characidium chupa (fig. 241)
7b. Pectoral rays eleven or fewer... ...8
8a. (7b) Eye larger, about 49% of head length; vertical bars thin,
more or less irregularly spaced and twelve or more in number...
Characidium sp. G (fig. 249)
8b. Eye smaller, 28-39% of head length; vertical bars wider and
usually eleven or fewer in number, better defined, more evenly
spaced... ...9

611
9a. (8b) Lateral line incomplete, with fewer than ten pored scales;
pectoral rays nine or fewer, usually only seven, the first two
extended as long filaments; dorsal-fin with a black spot near
anterior base; pectoral-fin with a black basal spot; body deeper,
its depth less than 3.6 in SL...
Characidium sp. H (no figure)
(Note: This is a rare, dwarf species, reaching only 20 mm SL; its
pigmentation is quite similar to Elachocharax spp.; because it was
recognized too late, it is not included in the species accounts)
9b. Lateral line complete, though sometimes difficult to see; pectoral
rays usually eleven or more, the first not filamentous; dorsal-fin
without spot at anterior base; pectoral-fin without black basal
spot; body more slender, its maximum depth four or more in SL...10
10a. (9b) Only 2-4 short fat gill rakers on lower limb of outer arch;
mouth subterminal, the gape located below a horizontal line drawn
through ventral margin of eye (fig. 234b); a few melanophores on
dorsal fin, some on membranes, most on outer tips of rays; no
concentration of pigment behind eye; at least some of the teeth
tricuspid; lateral scales 30-36; head length 15-18% of standard
length; interorbital width 47-55% of head width...
Characidium cf catenatum (fig. 247)
(Note: There are probably two very similar species keying here,
one with fewer lateral-line scales (30-33 vs 34-36), and a
shorter, wider head (HL 15% SL vs 18%, 10 55% HL vs 47%.)
10b. About eight thin elongate gill rakers on lower limb of outer arch;
mouth terminal, the gape in line with or above horizontal line
through ventral margin of eye (fig. 234c); dorsal fin pigmented,
with a bar through central portion and many spots along outer

612
third of rays; a concentration of dark pigment behind eye; teeth
conical, or with tiny basal lateral cusps; head length 24% SL; 10
37% HI Characidium cf zebra (fig. 237)
11a. (lb) Adipose fin absent; 16-17 interrupted vertical bars on body;
all premaxillary teeth conical... El achocharax .iunki (fig. 253)
lib. Adipose fin present; 8-10 wide vertical bars on body; some anteri¬
or premaxillary teeth tricuspid...
Elachocharax pulcher (fig. 255)
Species Accounts.
Ammocryptocharax elegans Weitzman & Kanazawa 1976
Voladorita Verde - South American Grass Darter
Fig. 235. Map: fig. 236. Couplet 2a.
Generic synonymy
Ammocryptocharax Weitzman & Kanazawa 1976 (type species: Ammocrypto¬
charax elegans. by original designation).
Specific synonymy
Ammocryptocharax elegans Weitzman & Kanazawa 1976: 325-346 (type
locality, Colombia, State of Meta, Vichada river drainage, Caño Muco
about 15 km west of Puerto Gaitán toward Puerto Carreño), seen.
Types. Holotype: at USNM. Paratypes FMNH. No specimen numbers
were given in the original description. Ibarra and Stewart (1987)
unfortunately did not include catalog numbers of paratypes.
Etymology. AMMOCRYPTOCHARAX is a combination of the names Ammo-
crypta, a genus of North American darters, and Charax a genus of South
American characid fishes, in reference to the superficial similarity
between Ammocryptocharax and Ammocrypta; ELEGANS = is from the Latin

613
Figure 235. Ammocryptocharax elegans.

614
meaning special, choice, fine, or select (Weitzman & Kanazawa 1976).
Description
Illustrations. Figs. 234a (head) & 235; Weitzman & Kanazawa
1976:332, fig 1., 334, fig. 2.
Di agnosis. The strictly inferior mouth is unique to this species
among Apure drainage characidiids. It is also the most elongate and
slender member of the group with a greatest body depth of only 12% SL,
versus almost always 17% or greater in the other species.
Size. It can reach at least 40 mm SL.
Morphology. The body is pencil-shaped and very elongate. The
head is conical and the snout is very pointed and elongated. The snout
length is greater than the horizontal diameter of the eye. The fins
are not as enlarged as in mos Characidiurn spp. The teeth in
Ammocryptocharax are all conical, and are arranged in only two rows on
the dentary (Weitzman & Kanazawa 1976).
Counts. DR 10-12, usually ii9; AR 9, usually ii7; CR 19, 17
branched; PR 7-10 usually ii-6-i; VR 5; LLS 39-41; TS 8, 4 above, 3
below lateral line; 12 circumpeduncular scales.
Measurements. See original description for details, the follow¬
ing measurements are from the Apure specimen: GBD/SL 0.12; EYE/HL 0.36;
HL/SL 0.17; IO/HL 0.31; CPD/SL 0.07 (mean % values).
Pigmentation. Although the single Apure drainage specimen cap¬
tured was bright grass green, I have collected two color phases from
the Guyana Shield populations green and mottled cryptic brown.
Distribution and Natural History
Range. It is known from the Orinoco Basin, the Guaporé River
along Brazilian and Bolivian border, and the main course of the Amazon
River in the state of Pará, Brazil (Weitzman & Kanazawa 1976).

Figure 236. Apure drainage distribution of Ammocryptocharax elegans.

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UNIVERSIDAD NACIONAL EXPERIMENTAL
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CUENCA DEL RIO APURE
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617
Apure distribution. Map: fig. 236. This species is known only from
the far southeastern corner of the drainage.
Abundance. RARE.
Number of specimens examined. 1.
Food. BENTHIC OMNIVORE. It probably feeds on aquatic insects.
Habitat. The specimen taken during this study was found in a black-
water stream in lowland savannas where there was abundant aquatic vegeta¬
tion and a moderate current. Green specimens are always found in bright
green grass (probably a sedge). They will change to a darker green, or
lose the green color entirely when kept in surroundings of colors other
than bright green. Brown specimens are always found among sticks and
twigs. Brown specimens placed in bright green grass did not change col¬
ors.
Importance. Ornamental.
Characidium cf zebra Eigenmann 1909
Voladorita or Majuca - South American Darter
Fig. 237. Map: fig. 238. Couplet 10b.
Generic synonymy
Characidium Reinhardt 1866:55 (type species: Characidium fasciatum
Reinhardt, by monotypy); Fowler 1950:253 (synonymy).
Chorimycterus Cope 1894:67 (type species: Chorimvcterus tenuis Cope, by
monotypy).
Nanognathus Boulenger 1895:3 (type species: Nanognathus borel1 ii
Boulenger, by monotypy).
Poecilosomatops Fowler 1906:323 (type species: Characidium etheostoma.
by original designation).

618
Figure 237. Characidium cf zebra.

619
Specific synonymy
Characidium zebra Eigenmann 1909:38 (type locality: Maripicru, a branch
of the Ireng, in the upper Rio Branco drainage, of northern Brazil,
near the Venezuelan border, seen.
Types. Holotype: FMNH 53547 (formerly CM 1151). Paratypes: CM
1152-IU 11687 (6); CM 1159-IU 11687 (14). Ibarra & Stewart (1987) report
that only seven paratypes remain at FMNH; CAS 60247(2) (IU 11680); CAS-SU
21775(1); 21910(1).
Etvmology. CHARACIDIUM = presumably refers to a diminutive form of
Charax.
Description
Illustrations. Fig. 237.
Diagnosis. This species is most similar to Characidium cf catenatum
but has more gill rakers on the lower limb of the outer arch (8 vs 2-3)
and has a strictly terminal mouth (subterminal in Ch. cf catenatum.
Size. It grows to about 50 mm SL.
Morphology. The body is long and slender and the mouth terminal.
The teeth are mostly conical, but sometimes have tiny, basal lateral
cusps.
Counts. DR 9-11; AR 7-9; CR 17-20; PR 7-10, usually 10; VR 7-9,
usually 9. (N=9).
Measurements. (N=9), given as percent of SL or HL as indicated:
GBD/SL 17-23 (21); EYE/HL 27-35 (30); HL/SL 23-27 (24); IO/HL 29-49 (37);
PDL/SL 39-48 (44); CPD/SL 10-13 (11).
Pigmentation. The basic pattern is a base color of light tan,
crossed with dark vertical bars. There is a small black spot at base of
the caudal fin.

Figure 238. Apure drainage distribution of Characidium cf zebra.


622
Distribution and Natural History
Range. It is known from Guyana, Venezuela and northern Brazil (Rio
Branco).
Apure distribution. Map: fig. 238. This species is most common in
northern Apure state, and in the Aguaro River system of the southeastern
corner of the drainage.
Habitat. It inhabits blackwater streams of the lowland savanna. It
can burrow into the sand to seek shelter.
Abundance. COMMON.
Number of specimens examined. 191 from 26 collections.
Food. BENTHIC OMNIVORE Its diet includes aquatic insects, espe¬
cially chironomid larvae, microcrustaceans and algae.
Importance. Ornamental.
Characidium sp. D
Voladorita Negra - Black South American Darter
Fig. 239. Map: fig. 240. Couplet 3a.
Comments. This species is similar to Characidium crandelli Steind-
achner 1917, described from near Boa Vista, in the R. Blanco drainage,
Brazil). It also resembles Ch. blennioides Eigenmann 1912, from Guyana.
Géry (1977) distinguished between the two "species groups," stating that
the Ch. crandel1i group has only the first ventral (pelvic-fin) ray un¬
branched, whereas the Ch. blennioides group has the first two unbranched
(at least on one side) and sometimes thickened.
Description
Illustrations. Fig. 239.
Diagnosis. This is the only dark Characidium in the Apure drain¬
age that is dark brown to black in overall pigmentation. In addition,

623

Figure 240. Apure drainage distribution of Characidium sp.


626
the caudal fin has distinct vertical bars and the first four pelvic-fin
rays are unbranched.
Size. It reaches about 40 mm SL.
Counts. DR 10-11; AR 7; CR 19; PR 10-13, usually 12; VR 8 (usually
only the first one unbranched); LLS 28; TS above LL 3, (N=5).
Measurements. GBD 0.17; EYE/HL 0.35; HL 0.19; I0/HL 0.44; PDL 0.45;
CPD 0.12. (N=5).
Pigmentation. It is very dark brown to black in life, with the tail
conspicuously barred.
Pistribution and Natural History
Range. It is known from the Amazon and Orinoco basins.
Apure distribution. Map: fig. 240. It has been taken only from
northern Apure State and the Aguaro River system in the southeastern
corner of the drainage.
Habitat. It lives in blackwater streams of the lowland savanna.
Abundance. UNCOMMON.
Number of specimens examined. 17 from 8 collections.
Food. Unknown, but probably a benthic omnivore.
Importance. Ornamental.
Characidium chupa Schultz 1944
Voladorita de Montaña - Andean Darter
Figs. 234b & 241. Map: fig. 242. Couplet 7a.
Specific svnonvmv
Characidium chupa chupa Schultz 1944b:284 (type locality for this
subspecies: upper Río Chama, Maracaibo Basin), seen; Mago L. 1970:74.
Characidium chupa torbesensis Schultz 1944b:286 (type locality for the
subspecies Ch. chupa torbesensis: Río Torbes, 1 km above Táriba,
Orinoco drainage), seen; Mago L. 1970:74.

Figure 241. Characidium chupa.

628
Types. For the subspecies Ch. chupa torbesensis Holotype: USNM
121415. Paratypes: USNM 121416 (7). For the subspecies Ch. chupa chupa
Holotype: USNM 121417. Paratypes: USNM 121418 (1), 121419 (80), 121420
(9).
Comments. Schultz (1944b) described both subspecies, Characidium
chupa chupa from the upper Río Chama in the Maracaibo Basin, and Ch. chupa
torbesensis from the Rio Torbes near San Cristobal in the upper Apure
drainage, Orinoco Basin, in the same paper.
Etymology. CHUPA = is a common name for this species in the Rio
Chama valley near Mérida.
Description
Illustrations. Figs. 234b & 241; Schultz 1944b:287, fig. 34.
Di agnosis. This species is characterized by a relatively high
pectoral-fin ray count of 12-15.
Size. This is a large Characidium. reaching at least 80 mm SL.
Morphology. It is a robust, heavy-bodied fish when adult.
Counts. DR 10; AR 7-9; CR 18-20; PR 10-13, usually 12-13; VR 8-9;
LLS 34-38; TS above LL 4-5. (N=5)
Measurements. GBD 0.21; EYE/HL 0.28; HL 0.17; IO/HL 0.44; PDL
0.43; CPD 0.12. (N=5).
Pigmentation. Schultz (1944b) gave the following description: The
color bars extend below the blackish lateral band and are not enlarged
into round blotches along the mid-axis as in C. chupa chupa. In life
this species is brightly colored, with yellow spots on head and gills
and yellow or orange on the fins.
Distribution and Natural History
Range. It is known from the Andes and Perijá mountains, of the
Orinoco and Maracaibo basins.

Figure 242. Apure drainage distribution of Characidium chupa.


631
Apure distribution. Map: fig. 242. In the Apure drainage this
fish is restricted to the piedmont and montane streams of the Andes.
Habitat. It is found in clear water with moderate to rapid current
over a rocky substrate in streams and rivers of the Andean piedmont and
mountains.
Abundance. RARE.
Number of specimens examined. 9 from 7 collections.
Food. BENTHIC OMNIVORE. Its diet includes aquatic insects,
especially chironomid larvae, microcrustaceans and algae (pers. com.
A1ex FIecker).
Reproduction. Though no field observations are available, the
bright colors of adults suggest some sort of courtship ritual.
Importance. Ornamental.
Characidium sp. K
Voladorita Transparente - Transparent South American Darter
Fig. 243. Map: fig. 244. Couplet 6a.
Comments. This species is similar to Characidium pel 1ucidum
Eigenmann 1909 from Guyana.
Description
111ustrations. Fig. 243.
Diaonosis. This species is characterized by the following
combination of characters: the middle of the side is marked with a
horizontal row of black spots; the gape is even with or above a
horizontal line from along ventral edge of eye (fig. 1C); there are
usually only seven pectoral-fin rays; the greatest body depth is about
20% SL; the head is relatively long, its length about 17% SL; the
interorbital width is about 40% HL.

632
Figure 243. Characidium sp. "K".

Figure 244. Apure drainage distribution of Characidium sp. K.

CUENCA DEL RIO APURE
(«CALA UATCI
too
cr>
co
-p*

635
Size. This is a dwarf Characidium that seldom exceeds 40 mm SL.
Morphology. It is an elongate and slender-bodied Characidium. the
head longer than in most species. The pectoral fins reach the ventrals,
and the outer pectoral rays are not thickened.
Counts. DR 10-11; AR 7; CR 16-17; PR 7; VR 9; LS 30; TS above LL 4;
(N =8).
Measurements. GBD 0.20; EYE/HL 0.34; HL 0.25; IO/HL 0.30; PDL 0.37;
CPD 0.10.
Pigmentation. The body is transparent in life, but in preservative
numerous tiny chromatophores give it a dusky tan background color.
There is a row of irregular black spots down the middle of the sides.
Distribution and Natural History
Range. It is known from the Orinoco Basin.
Apure distribution. Map: fig. 244. Collections are all from the
high llanos.
Habitat. Few specimens have been collected, but this species has
been found so far in Whitewater streams with slow to moderate currents.
Abundance. RARE.
Number of specimens examined. 36 from 4 collections.
Food. Unknown but probably a benthic omnivore.
Importance. Ornamental.
Characidium sp. J
Voladorita Arenera - South American Sand Darter
Fig. 245. Map: fig. 246. Couplet 6b.
Comments. This species is similar to Characidium pteroides
Eigenmann 1909, from Guyana.

636
Figure 245. Characidium sp. "J".

637
Description
Illustrations. Fig. 245.
Diagnosis. This Characidium is characterized by its small size,
an incomplete lateral line (but its scales are deciduous and usually
lost), and a pigmentation pattern of a light background with light
brown crescents on the sides. It lacks the dark vertical bars typical
of most Apure drainage characidiids.
Size. This is a very small, dwarf Characidium that seldom exceeds
30 mm SL.
Morphology. It is elongate and slender, and well adapted to burrow¬
ing into the sand.
Counts. DR 8-11; AR 6-7; CR 15-19; PR 8-10; VR 6-9; LS 25-29; TS
above LL 4. (N=7).
Measurements. GBD 0.16; EYE/HL 0.39; HL 0.17; I0/HL 0.40; PDL
0.41; CPD 0.11 (mean values), (N=7).
Pigmentation. The body is light tan with irregularly spaced light
brown crescents along the sides.
Distribution and Natural Hi story
Range. It occurs in Guyana and the Orinoco Basin.
Apure distribution. Map: fig. 246. It has been taken only from
the Aguaro River system.
Habitat. It lives in blackwater lowland savanna streams, usually
over white sand substrates.
Abundance. RARE.
Number of specimens examined. 24 from 6 collections.
Food. Unknown, but it is probably a benthic omnivore.
Importance. Ornamental.

Figure 246. Apure drainage distribution of Characidium sp. J.

LO
VO

Characidium cf catenatum Eigenmann 1909
Voladorita - South American Darter
640
Fig. 247. Map: fig. 248. Couplet 10a.
Specific synonymy
Characidium catenatum Eigenmann 1909:40 (type locality: Warraputa,
Rockstone, Crab Falls, Guyana); Eigenmann 1920:10 (from Rio Guaire
near Caracas and Rio Tuy in Concejo, Venez.), seen; Schultz 1944b:279.
Types. Holotype: FMNH 52761 (CM 1153). Paratypes: of the original
paratypes CM 1154-III 11681 (12 specimens) and CM 1155-IU 11682 (2 specs.)
only 6 remain at FMNH (Ibarra & Stewart 1987).
Comments. There may be two species included here. The lateral-line
scale counts seem to fall into two groups, and the distribution map indi¬
cates the possibility of separate highland and lowland populations.
Etymology. CATENATUM = chain, from Latin, referring to the chain¬
like pattern of the bars on the upper sides of this species.
Description
Illustrations. Fig. 247; Eigenmann 1912: pi. 38 (5-6).
Diagnosis. The subterminal mouth and low gill raker count (2-3)
separate this species from other similarly pigmented Characidium in the
Apure drainage.
Size. This is a medium-sized Characidium, reaching 45 mm SL.
Morphology. The mouth is distinctly subterminal.
Counts. DR 9-11; AR 6-8; CR 17-19; PR 8-11; VR 7-9; LLS 30-33 in
some populations, 34-36 in others; TS above LL 4. (N=12).
Measurements, given as % SL or HL as indicated: GBD/SL 19; EYE/HL
33; HL/SL 17; IO/HL 55 in some populations, 47 in others; PDL/SL 44;
CPD/SL 12. (N=12).

Figure 247. Characidium cf catenatum.

Figure 248. Apure drainage distribution of Characidium cf catenatum.

MUit •a«»ic4
643

644
Pigmentation. The body is whitish to light brown, with 9-10 black
or brown vertical bars that are interrupted on midside. In addition to
those bars, the sides have 4-5 horizontal rows of faint dots (one per
scale). There is a dark black spot at the base of the caudal fin. A brown
line extends from the eye to the tip of the snout.
Distribution and Natural History
Apure distribution. Map: fig. 248. There seem to be two separate
populations, one in the piedmont and the other in low llanos. This sepa¬
ration could account for the different counts noted above, and is another
indication that two species may be involved.
Habitat. It is known from a wide variety of lotic habitats, in
both the piedmont and llanos.
Abundance. ABUNDANT.
Number of specimens examined. 389 from 82 collections.
Food. BENTHIC OMNIVORE. The diet includes aquatic insects
(especially chironomid larvae), microcrustaceans and algae.
Importance. Ornamental.
Characidium sp. G
Voladorita - South American Darter
Fig. 249. Map: fig. 250. Couplet 8a.
Description
Illustrations. Fig. 249.
Diagnosis. Of all the Apure drainage Characidium species, this one
has the shortest, deepest body. The eye is particularly large, its
diameter comprises 49% of the head length. The pigmentation pattern is
also distinctive, consisting of numerous thin vertical bars (more than
in any other Apure drainage species).

645
Figure 249. Characidium sp. "G".

646
Size. This small species seldom exceeds 30 mm SL.
Morphology. The body is short, high, and stocky. The head is wide,
with a bluntly rounded snout.
Counts. DR 8-10; AR 7-9; CR 19; PR 9; VR 7-8; LLS 31-33; TS above
LL 4, (N=7).
Measurements, given as % SL or HL as indicated: GBD/SL 22; EYE/HL
49; HL/SL 16; IO/HL 55; PDL/SL 43; CPD/SL 13 (mean values), (N=7).
Pigmentation. The base color of the body is light brown. There are
11-15 thin vertical bars on the sides, some of them double, and often
united across the dorsum. Many specimens have additional thin, short,
incomplete vertical bars in between the wider ones. The lateral midline
has a dark stripe that extends from the tip of the snout back through the
eye and onto the opercle. The base of the caudal fin is marked with a
small black dot. A black line extends forward along the ventral midline
from in front of the anal fin.
Pistribution and Natural History
Apure distribution. Map: fig. 250. It is widespread throughout
the llanos and piedmont.
Habitat. It lives in a wide variety of lotic habitats.
Abundance. ABUNDANT. This is the most common characidiid in the
Apure drainage.
Number of specimens examined. 464 from 83 collections.
Food. BENTHIC OMNIVORE. Its diet includes aquatic insects,
microcrustaceans and algae. I have observed this species in aquaria,
hovering just beneath floating aquatic vegetation, staring intently at
the surface for several minutes, then making a quick jab at some small
(invisible to me) aquatic creature.
Importance. Ornamental.

Figure 250. Apure drainage distribution of Characidium sp. G.


649
Characidium voladorita Schultz 1944
Voladorita - South American Darter
Fig. 251. Map: fig. 252. Couplet 4a.
Specific synonymy
Characidium voladorita Schultz 1944b:280, fig. 32 (type locality: 4 km
above Motatán in the Río Motatán, Maracaibo Basin), seen; Mago L.
1970:74.
Types. Holotype: USNM 121407. Paratypes: USNM 121408 (40); 121409
(15); 121410 (27); 121411 (3); 121412 (10); 121413 (26); 121414 (96).
Comments. This species was originally described and hitherto known
only from the Maracaibo Basin. I have not carefully compared Apure speci¬
mens with Maracaibo specimens, but the presence of Characidium species on
both sides of the Andes is not unprecedented, as for example in the case
of Ch. chupa
Etymology. VOLADORITA = the common name of this species in Spanish,
meaning "little flyer." The name "volador" is given to Lebiasina ervthri-
noides a species frequently sympatric with Characidium in the Andes. The
name "voladorita" means little volador, and is probably a result of the
common but erroneous belief of rural people that all small fishes must be
juveniles of some larger fish they know and use. Thus when Schultz asked
for the name of this Characidium, they gave the diminutive form of the
name for Lebiasina.
Description
Illustrations. Fig. 251; Schultz 1944b:280, fig. 32.
Diagnosis. This is the only species of Characidium in the Apure
drainage with a naked area between the pectoral fins.
Size. It reaches about 45 mm SL.

650
Figure 251. Characidium voladorita.

651
Counts. Schultz (1944b) gave the following for the Maracaibo
population: DR i ii8-i ii9; AR i ii6; PR i ii8-i ii9; VR i7; GR 5+10 on first
arch; LLS 30-33; TS 3 1/2 above, 2 1/2 - 3 below; 10-12 predorsal
scales. Apure drainage specimens usually fall into these ranges.
Measurements. GBD 22% SL; EYE 33% HL; HL 16% SL; 10 43% HL; PDL 45
% SL; CPD 13% SL (mean values).
Pigmentation. The live coloration is quite striking in the
breeding season, when the males become brick red on the entire ventrum,
rear portion of the body and tail. The dorsal fin is red and yellow,
and the head is bordered posteriorly with a yellow band. In alcohol it
is tan with darker vertical bars when young, but adults can lose the
vertical bars almost completely. There is always at least a trace of a
humeral spot just behind the opercle.
Pistribution and Natural History
Range. It is known from the Maracaibo Basin, as well as in
adjacent areas of the Andean piedmont in the Apure drainage. It
probably also extends into the Colombian Andes.
Apure distribution. Map: fig. 252. This is the first record of
this species in the Orinoco Basin. It is restricted to Andean streams
and rivers.
Habitat. It occurs in Andean piedmont and montane streams with
fast currents and rocky substrates.
Abundance. COMMON.
Number of specimens examined. 302 from 46 collections.
Food. BENTHIC OMNIVORE. It feeds on aquatic insects, especially
chironomid larvae, microcrustaceans and algae (Alex Flecker, pers. com.).
Importance. Ornamental.

Figure 252. Apure drainage distribution of Characidium voladorita.

653

654
Elachocharax cf .iunki Géry 1971
Voladorita Enana - Dwarf South American Darter
Fig. 253. Map: fig. 254. Couplet 11a.
Generic synonymy
Elachocharax Myers 1927:114 (type species: Elachocharax pulcher Myers
1927), by monotypy; Géry 1977:122; Weitzman & Kanazawa 1978:160
(redefinition); Weitzman & Géry 1981:888 (revision).
Elacocharax Weitzman & Kanazawa 1976:328 (misspelling).
Geisieria Géry 1971:154 (type species Gei1eria iunki 1971), by
monotypy; Gery 1977:122.
Soecific synonymy
Geisleria .iunki Géry 1971:154 (type locality: Brazil, Rondonia, Río Novo,
tributary to Rio Jamari about 45 km east of Porto Velho, Rio Madeira
basin; Géry 1977:122; (listed, brief comments); Weitzman & Kanazawa
1978:168 (relationships); Weitzman & Géry 1981:887 (key; redescription);
Weitzman 1986:740 (key).
Types. Holotype: collection of J. Géry. Paratypes: MZUSP 14008 (1);
and four specimens in the collection of J. Géry.
Etymology. ELACHO = insignificant, from the Greek, and CHARAX a genus
of Characidae; JUNKI = to honor Dr. W. Junk of INPA in Manaus, Brazil.
Description
Illustrations. Fig. 253.
Diagnosis. This species is distinguished from all Characidium species
by the possession of more than 16 dorsal-fin rays. It differs from Elacho¬
charax pulcher by the lack of an adipose fin and the absence of tricuspid
teeth on the premaxilla. Pigmentary features vary too much to be useful as
diagnostic characters.
Size. It is a dwarf species seldom exceeding 20 mm SL.

655
Figure 253. Elachocharax .iunki

656
Morphology. This species has a short compressed body. The lateral
line is incomplete. The anal-fin origin is at or slightly posterior to a
vertical line drawn through the posterior termination of the dorsal -fin base
(Weitzman 1986). The teeth in both jaws are all conical.
Counts. Teeth on ectopterygoid 4-5, on inner row of dentary 13-16;
upper limb gill rakers 3; vertebrae 30-33; scales rows between anterior base
of dorsal fin and pelvic fin 8; lateral scales 27-33 Weitzman 1986).
Measurements. Snout length about 6-8% SL. Caudal peduncle length
about 16-19% SL (Weitzman 1986).
Pigmentation. Although this species is reported (Weitzman 1986) to
have 13 to 14 narrow vertical dark bars in the caudal fin, most Apure drain¬
age specimens do not show this pattern. The body is dark and crossed with
vertical bars. The anal fin is marked with two strong black pigment bars,
whereas the pelvic fin has only one.
Distribution and Natural History
Range. This species occurs in the upper Rio Negro drainage in Brazil
and Venezuela, in the Casiquiare Canal area, and in the upper Orinoco Basin
in Venezuela (Weitzman 1986).
Apure distribution. Map: fig. 254. It is known only from the Aguaro
River system.
Habitat. It lives in blackwater streams in the low llanos, usually in
areas with aquatic vegetation, or with finely divided roots from terrestrial
vegetation growing along the banks.
Abundance. RARE.
Number of Specimens Examined. 7 from 1 collection.
Food. BENTHIC OMNIVORE.
Importance. Ornamental.

Figure 254. Apure drainage distribution of Elachocharax junki.

10*
9*
»•
I*
CUENCA DEL RIO APURE
658

Elachocharax pulcher Myers 1927
Voladorita Enana - Dwarf South American Darter
659
Fig. 255. Map: fig. 256. Couplet lib.
Specific synonymy
Elachocharax pulcher Myers 1927:114 (type locality: Venezuela: Caño de
Quiribana, near Caicara,), seen; Mago L. 1970:74; Géry 1977:114, 119.
Description
Types. Holotype: IU 17676. Paratype: IU17677
Illustrations. Fig. 255; Géry 1977:118 (of holotype).
Di agnosis. see key. The presence of the adipose fin distinguish¬
es this species from E. .iunki the only other species known from the
Apure drainage. The high number of dorsal fin rays is sufficient to
distinguish from species of Characidium.
Size. This dwarf species seldom exceeds 20 mm SL, and is usually
taken much smaller.
Morphology. The lateral line is incomplete, with only 5-6 pored
scales. The mouth is small, with very little gape, but the maxilla
reaches a point vertical of the anterior border of the eye. Some of
the anterior premaxillary teeth are bi- to tricuspid. The body is much
compressed, the head is especially narrow anteriorly with a pointed
snout.
Counts. DR 18; AR 8; LS 25, TS 6 from dorsal to pelvics (this
count was given as 8 by Weitzman 1986, this difference might be caused
by different counting methods, or the Apure specimens might represent a
different taxon).
Measurements. The head length is equal to the body depth and
measures 3 1/2 times in the SL.

660
Figure 255. Elachocharax pulcher.

661
Pigmentation. This tiny fish is usually very darkly pigmented in
life, with horizontal zig-zag lines between scale rows on sides. It
body is crossed by eight diffuse but wide dark bands. The caudal fin
is sometimes faintly banded. The dorsal fin has two dark stripes one
very near its base, and the other located distally, but not along the
outer margin of the fin. The eye is crossed by two lines, one horizon¬
tal from the chin, and the other vertical from the occiput. This color
pattern undoubtedly helps this species blend in the roots and other
vegetation that form its natural refuge.
Etymology. PULCHER = from the Latin for pretty.
Distribution and Natural History
Range. This species is probably present in most of the upper Rio
Negro in Brazil, Colombia and southern Venezuela, as well as isolated
localities throughout the Orinoco Basin (in appropriate blackwater
habitat). As with most characiformes, a closer examination of the
isolated populations present in the Apure River drainage might reveal
taxonomic differences.
Apure distribution. Map: fig. 256. It is known only from the
Aguaro River system.
Habitat. It is found in blackwater streams of the low savanna.
This species is said to live among fine roots of plants growing along
the banks (Géry 1977), and is almost always associated with some sort
of finely divided cover.
Abundance. RARE.
Number of specimens examined. 8 from 2 collections.
Food. BENTHIC OMNIVORE.
Importance. Ornamental.

Figure 256. Apure drainage distribution of Elachocharax pulcher.

663

664
Chilodontidae
The Chilodontidae (also spelled Chilodidae by Géry [1977]) is a
small family of but two genera and three or four species (Géry 1977).
So far, only one of these, Chilodus punctatus, has been found in the Apure
drainage. Although this group is easily recognized and
distinguished, its taxonomic rank is still unclear. For example Nelson
(1984) included the "Chilodontinae" as a subfamily of the Curimatidae,
but Vari (1989) continues to recognize it as a distinct family. Vari
(1983) considered the anatomy of the included species’ epibranchial
organ and pectoral girdle is sufficiently distinct to warrant familial
recognition. They are most closely related to the Prochi 1odontidae and
together with that family they form a sister group to the Anostomidae-
Curimatidae pair (Vari 1983).
The chilodontids are the popular "headstanders" of the aquarium
trade. They swim head-down usually in quiet waters amidst aquatic
vegetation, feeding on aquatic insect larvae and other small creatures.
Little is known about their natural history.
Species Accounts.
Chilodus punctatus Mül1er & Troschel 1844
Spotted Headstander - Conchuo
Fig. 257. Map: fig. 258.
Generic synonymy
Chilodus Milller & Troschel 1844:85 (type species: Chi 1 odus punctatus
Müller & Troschel, by monotypy; [not Chilodus Giebel 1847]); Eigenmann
1912:273; Fowler 1950:214 (synonymy).

665
Figure 257. Chilodus punctatus.

666
Specific synonymy
Chi 1 odus punctatus Miiller & Troschel 1844:86 pi. 4, fig. 2 (type
locality: Lake Amucu, Guyana), seen; Eigenmann 1912:273 (synonymy,
description); Géry 1977:214.
Citharinus chilodus Valenciennes in Cuvier & Valenciennes 1849:103.
Caenotropus punctatus Gunther 1864:297.
Types. Lectotype: ZMB 23599. Paralectotype: ZMB 24074.
Comments. Caenotropus 1abyrinthicus occupies neighboring drain¬
ages to the south. It’s ventral mouth makes it easy to distinguish
from this species.
Etymology. CHIL = lips, ODUS = teeth; PUNCTATUS = spotted.
Description
Illustrations. Eigenmann 1912:pi. 35, fig. 4; Géry 1977:212, 213;
Román 1983:95.
Di agnosis. Chilodus punctatus can be recognized at once by the
distinctive body form and pigmentation (fig. 257). There is a black
lateral stripe from the tip of the snout, through the eye and back to
the base of the tail. The tips of the first few dorsal rays are black.
Each scale has a dark spot over a whitish to tan background. This
species is somewhat similar to curimatids and anostomids, but is easily
distinguished by the presence of a single row of feeble, truncate teeth
in each jaw, and by the unique, dilated fourth gill arch.
C. punctatus can be distinguished from C. grácil is by its deeper
body (GBD 2.7-3.2 vs 3.3-3.6 in C. grácil is). C. zunevi always lacks a
lateral stripe, has spots around the pectoral fin, and has dorsal and
ventrally placed spots of about equal intensity. In C. punctatus the
lateral stripe is usually present although it is sometimes zigzag, the

667
spots are usually darker dorsally and are absent around the pectoral
fin (Isbriicker & Nijssen 1988).
Size. It usually does not exceed 70 mm SL.
Morphology. The body is compressed and fairly deep with an
elevated back. The head is conical, with a small, terminal to upturned
mouth. The scales are large and entire. The lateral line is straight
and complete.
Counts. DR 11; AR 12; LLS 25-27; TS 8-9.
Measurements. HL 3.7 in SL; GBD 3 in SL; EYE .75 in snout, 3 in
HL, equal to interorbital (Eigenmann 1912).
Pigmentation. The base color is tan to silvery. A black lateral
stripe (sometimes expressed as an interrupted series of dots and
blotches) extends from the tip of the snout to the base of the caudal
fin; each scale of the sides and dorsum is marked with a large black
spot, and the black pigment sometimes "runs together" to form a net-
like pattern. The dorsal fin has a large black blotch near its tip,
and several smaller black spots proximally. The anal fin is dark, but
the other fins are colorless. The belly and throat are silvery white.
Géry (1977) noted that the usual spotted pigmentation pattern disap¬
pears during spawning and is replaced by a broad humeral spot.
Distribution and Natural History
Range. It is known from British Guiana, and the Orinoco and
Amazon basins.
Apure distribution. Map: fig. 258. It has been found only in
northern Apure state.
Habitat. It inhabits blackwater creeks, bajios and esteros.
Abundance. UNCOMMON.

Figure 258. Apure drainage distribution of Chi 1odus punctatus.

CUENCA OEL RIO APURE
669

670
Number of specimens examined. 156 from 24 collections.
Food. OMNIVORE. It probably feeds mainly on minute animal life
such as microcrustaceans, aquatic insects, and other aquatic inverte¬
brates that live on aquatic vegetation, as well as vegetable matter.
Reproduction. Strategy probably rl. Eggs are scattered in
aquatic vegetation or in aquaria on artificial nylon spawning mops.
The male pushes the female into the substrate then pushes its caudal
peduncle under that of the female, at which time spawning occurs. Eggs
are large, swelling to 2 mm, and hatch in 3-4 days (Sterba 1972).
Importance. Ornamental.
Ctenoluciidae
The pike characins are elongate, barracuda-like piscivores that
prowl just beneath the water’s surface seeking their prey. The family
is not a large one, and Géry (1977) cited only four known species in
two genera, Ctenolucius and Boulengerel 1 a. The species are readily
recognized by their elongate, slender jaws and numerous, conical teeth
present in both the upper and the included lower jaw, as well as on the
ectopterygoid bones. The teeth point backward toward the inside of the
mouth to prevent the escape of their prey. Either the tip of the snout
or the tip of the lower jaw, bears a fleshy appendage that is a charac¬
teristic feature of this group. Boulengerel1 a lucius, is the only
species that has been found so far in the Apure drainage, but the other
three species in the family are present in other parts of Venezuela.
Ctenolucius hujeta occurs in the Maracaibo Basin, and Boulengerella
lateristriga and ]L maculata are present in the Amazonas Territory and
Bolivar state.

671
Species Accounts.
Boulengerel1 a lucius (Cuvier) 1817
Pike Characin - Agujeta
Fig. 259. Map: fig. 260.
Generic synonymy
Boulengerella Eigenmann 1903:147 (type species: Xiphostoma lateristriga
Boulenger, by original designation).
Xiphostoma (not Kirby & Spence 1828 in Hemiptera) Spix i_n Agassiz
1829:60 (type species: Xiphostoma cuvieri Spix j_n Agassiz, by
monotypy; Valenciennes in Cuvier & Valenciennes 1849:X111; Jordan &
Evermann 1917:132
Xiphostomus Swainson 1839:290 (type species: Xiphostomus cuvieri Spix).
Specific synonymy
Hydrocynus 1ucius Cuvier 1817:168 (type locality: rivers of Brazil),
seen.
Hydrocyon lucius Cuvier 1819:359 (type locality: Brazil).
Xiphostoma lucius Miiller & Troschel 1844:94 (locality not indicated).
Xi phostoma cuvieri Agassiz i_n Spix 1829:78, pi. 42 (type locality:
rivers of Brazil); Pellegrin 1899:157 (Apure River, Venezuela).
Hvdrocvnus cuvieri Eigenmann 1910:446.
Boulengerella cuvieri Schultz 1944b:261; Fowler 1950:328 (synonymy).
Xi phostoma ocel1atum Schomburgk 1841:245, pi. 23. (type locality:
Essequibo; Negro and Branco rivers, Brazil).
Xiphvstoma (Terror) ocel1atum 1895:296.
Hvdrocvnus ocellatus Eigenmann 1910:446 (alternative spelling).
Xiphostoma oservi Castelnau 1855:76, pi. 40. fig. 1 (type locality:
Tocantins, Brazil).

^rr*sr 11
Figure 259. Boulengerella lucius.

673
?Xiphostoma longipinne Steindachner 1876:132, (type locality: mouth of
Rio Negro).
Boulengerella lucius Mago L. 1970:74.
Boulengerella (Spixostoma) lucia Géry 1977:106 (key, synonymy).
Comments. Géry (1977) called this species B. lucia, changing the
ending of the specific name to make it agree with the genus. I have
retained the original spelling (B. lucius). He placed it in the mono-
typic subgenus Spixostoma, and listed B. ocel1atus, B. cuvieri and B.
longipinne as probable synonyms. He regarded the other subgenus,
Boulengerella, as comprising B_^ 1 ateristriga and IL. maculata, both of
which have an incomplete lateral line, and fewer lateral-line scales
(78-92).
Etymology. BOULENGERELLA = so named to honor George Albert Bou-
lenger, a Belgian ichthyologist, who worked at the British Museum of
Natural History for many years; LUCIUS = refers to the genus of the
similarly shaped, and ecologically parallel species of North American
pi ke.
Description
Illustrations. Fig. 259; Géry 1977:107.
Diagnosis. The elongate, canine-studded jaws, and fleshy append¬
age on the tip of the upper jaw clearly distinguish this fish from
superficially similar species. It has a complete lateral line, with
97-110 scales in the lateral series above or below it. The other mem¬
bers of this genus, B. lateristriga and B. maculata have an incomplete
lateral line, and fewer scales (78-92) in the lateral series.
Size. It is said to reach one meter, but is usually seen at less
than half that size.

674
Morphology. The body is somewhat terete, elongate and slender.
The jaws are also very elongate and narrow, and beaklike. The dorsal
fin is placed far back on the body, posterior to the pelvic fins. An
adipose fin is present. The tail is large, and the lower lobe is
usually the larger. The lateral line is complete. The scales are
finely ctenoid.
Counts. DR 10; AR 10-11; PR 25-30; VR 8; LLS 97-110.
Measurements. HL 2.8-3.4; GBD 6.5-7.5 (both in SL); eye 4.3-5 in
snout, 8-8.5 in HL, 1.5-1.75 in 10 (Eigenmann 1912).
Pigmentation. The body is silvery to golden in life with a few
scattered spots situated dorsally in some specimens. Young specimens
have a wide, black, lateral stripe that extends from the base of the
caudal fin forward through the eye and out to the tip of the fleshy
appendage on the snout. The caudal fin is marked with a black spot or
ocellus at the base, its outermost rays are white.
Pistribution and Natural Hi story
Range. It occurs in Guyana and other northern coastal drainages
of South America and in the Amazon Basin.
Apure distribution. Map: fig. 260. In the Apure drainage this
species has been found only from northern Apure state between Bruzual
and Mantecal, with the exception of one locality in the lower Suripá
river.
Habitat. It is a lowland species often found near the quiet,
vegetated edges of flowing waters of the low llanos.
Abundance. RARE.
Number of specimens examined. 11 from 10 collections.

Figure 260.
Apure drainage distribution of Boulengerella lucius.

676

677
Food. CARNIVORE. It is almost exclusively piscivorous, but
probably catch other prey on occasion. It feeds near the water’s
surface, frequently taking refuge behind cover near the faster moving
areas of streams. From there it darts out to take prey that are strug¬
gling against the current.
Importance. Ornamental. This species is also of some value as a
sport fish, but because of the numerous intermuscular bones is not
highly valued as a food fish.
Curimatidae
Many curimatids are analogous in general body plan, feeding strate¬
gies and ecological niches to the carps and suckers of North America.
They are toothless as adults, but juveniles of some species have teeth
until they reach a SL of some 20 mm, after which the teeth are reab¬
sorbed. Almost all are benthic detritivores that feed on algae, dia¬
toms, protozoans and bacteria from the surface layers of mud. They
abound in almost all water bodies, and figure strongly in the biomass
pyramid of llaneran lagoons and ponds, but also are common in slower
sections of streams and rivers. A few have adopted a more pelagic,
perhaps mainly pianktivorous life style, as is evidenced by their keeled
abdomen and modified gill rakers. Those species are usually found in or
near the main channels of large rivers. All have a gut that is quite
long, as in herbivores in general. The vast majority of the curimatids
are plain colored, silvery fishes (though often with a black spot on the
dorsal fin or caudal peduncle), with only small deviations from their
generalized body plan. They are mainly diurnal, and have large eyes,
often partly covered by a protective membrane.

678
Vari (1983, 1989) summarized and commented on the extremely
varied phylogenetic schemes proposed in the past for Curimatidae, and
established their relationships with the Prochilodontidae, Anostomidae
and Chilodontidae. He concluded that each of these four families is
monophyletic and that together they constitute a monophyletic assem¬
blage, with two pairs of sister groups: Curimatidae with Prochilodonti¬
dae and Anostomidae with Chilodontidae.
Fernández Y. (1948) enthusiastically, and for the most part
erroneously, split the family into 27 genera, and estimated some 109
species in this group. Géry (1977) estimated the family to contain 110
species and Vari (1983) estimated 120. Mago L. (1970) listed only nine
species for Venezuela, but many additional species have been discovered
and described since then.
Most recently, Vari (1989) listed 128 nominal taxa, but mentioned
"about 100" as the number of valid species in his phylogenetic study of
the family. He recognized only eight genera: Curimatopsis which in¬
cludes dwarf species with an incomplete lateral line, and basal spurs
on the middle caudal-fin rays of males; Potamorhina which includes
large, riverine species with small scales (85-110 LLS), numerous anal-
fin rays (13-19), and enlarged opercular bones; Curimata, which are
large species with numerous flaps and folds on the roof of the mouth;
Psectroqaster, relatively large species characterized mainly by spe¬
cializations of the epibranchial bones; Steindachnerina the largest
genus, and one that contains many of the smaller species formerly in
Curimata; Pseudocurimata, a genus present only west of the Andes;
Curimatella. comprised of small to medium sized species with scaled
caudal fins; and Cyphocharax. an assemblage of "leftover species" for

679
which no synapomorphies were discernible, and which includes three
phenetically recognizable groups: small species with a caudal peduncle
spot; small species from southern South America with an incomplete
lateral line; and a third group that includes the distinctively pig¬
mented curimatids. I have followed Vari (1989) for all generic synony¬
mies, and for the correct spellings and dates of species.
The names of the eight Apure drainage "species" employed here are
still tentative. They were determined using Vari’s preliminary, unpub¬
lished key to Venezuelan species (1983, photocopy) and Géry (1977). Since
almost all of the species are poorly described, many are sympatric, and
most are not easily recognized upon superficial visual inspection, I have
opted to "lump" whenever possible. It is therefore likely that I have not
recognized some closely related species.
Key to the Apure Drainage Species of Curimatidae.
la. Caudal fin densely scaled almost to the tips of the rays...
(genus Curimatella. 2 spp.)... ...2
lb. Caudal fin at most scaled only near base... ...3
2a. (la) Base of caudal fin with distinct black spot, separate from
lateral stripe... Curimatella bolivarensis (fig. 263)
2b. Base of caudal fin without a distinct black spot, separate from
lateral stripe... Curimatella immaculata (fig. 265)
3a. (lb) Lateral-line scales 45 or more from upper junction of
opercle to base of caudal fin... ...4
3b. Lateral-line scales 44 or fewer... ...7
4a. (3a) Lateral-line scales more than 90; transverse scales (from
dorsal-fin to pelvic-fin insertions) more than 40...
Potamorhina altamazonica (fig. 269)

680
4b. Lateral-line scales 45-60; transverse scales 15-25... ...5
5a. (4b) Mouth inferior... Steindachnerina sp. (fig. 273)
5b. Mouth terminal... ...6
6a. (5b) Interorbital width 53-58% of head length; gill rakers long
and slender, about 40 on first arch; AR iiil0-iiil2; ventrum,
pectoral, pelvic, and anal fins usually red in life...
Curimata cerasina (fig. 261)
6b. Interorbital width less than 52% of head length; gill rakers
replaced by membranous flap (actually reduced to tiny protuber¬
ances along base of flap); anal rays iii8-iii9; ventrum and
fins not red... Psectroqaster ciliatus (fig. 271)
7a. (3b) Dorsal fin with black dot, often faint or faded; lateral
scales not usually outlined in black...
Steindachnerina argéntea (fig. 275)
7b. Dorsal fin plain, without black dot; lateral scales often
outlined with black... Cyphocharax spilurus (fig. 267)
Species Accounts.
Curimata cerasina Vari 1983
Cherry Curimatid - Coporo Pechirojo
Fig. 261. Map: fig. 262. Couplet 6a.
Generic synonymy
Curimata Bose 1817 (type species: Salmo edentulosus Bloch 1794, by
monotypy).
Curimatus Valenciennes in. Cuvier & Valenciennes 1849:7 (alternative
spelling).

681
Figure 261. Curimata cerasina.

682
Semitapicis Eigenmann & Eigenmann 1889b (type species: Charax
planirostris Gray, 1854, by original designation).
Peltapleura Fowler 1906:300 (type species: Salmo cyprinoides Linnaeus,
by original designation).
Acuticurimata Fowler 1941 (type species: Curimatus macrops Eigenmann &
Eigenmann 1889b, by original designation)
Allenina Fernández Y. 1948 (type species: Curimata murieli Allen 1942 in
Eigenmann & Allen 1942, by original designation).
Lambepiedra Fernández Y. 1948 (type species: Lambepiedra al 1eni
Fernández Y. 1948, by original designation).
Bi tricari nata Fernández Y. 1948 (type species: Curimatus schomburqkii
Giinther 1864, by original designation).
Bondichthys Whitley 1953 (type species: Curimatus mivartii Steindachner
1878, by original designation).
Stupens Whitley 1954 (type species: Curimatus simulatus Eigenmann &
Eigenmann 1889b, by original designation).
Semitapiscis Braga & Azpelicueta 1983 (unjustified emendation of
Semitapicis).
Specific synonymy
Curimata cerasina Vari 1983:27 (type locality: Venez., Apure state,
flooded area along road from San Fernando de Apure to Arichuna, 15 km
SW of San Fernando de Apure), seen.
Curimata schomburqkii Mago L. 1967:254; Román 1985:135 (photo,
description).
Curimata ciprinoides Taphorn & Lilyestrom 1984:55 (biology).
Types. Holotype: MBUCV V-140.25. Paratypes: at MBUCV, USNM,
CAS, FMNH, BMNH, ANSP, AMNH, MCZ.

683
Comments. The name Curimata instead of Curimatus is used in
compliance with Opinion 722 (1966) of the International Commission of
Zoological Nomenclature.
Etymology. CURIMATA = a native (Brazilian) name for these fish¬
es; CERASINA = cherry like, referring to the red ventrum.
Description
Illustrations. Fig. 261. Vari 1983:40; Román 1985:135 (as C.
schomburqki i).
Di agnosis. This is the only large red-bellied curimatid in the
Apure drainage. It is further distinguished by the lack of scales on
the caudal fin, except at the base; by a very wide head (the interor¬
bital width is 53-58% of the head length); the presence of about 40
elongate slender gill rakers on the first gill arch; and 42-50 lateral¬
line scales.
Size. This is a large curimatid that grows to about 170 mm SL.
Morpholoqy. The scales are large and easily visible without
magnification. The chest is flattened and has distinct lateral longi¬
tudinal keels from level of a vertical through the pectoral-fin inser¬
tion to the insertion of the pelvic fin. There is a well developed
median keel posterior to the pelvic-fin insertion (Vari 1983). The
adipose fin is large. The mouth is terminal, with large lips.
Counts. DR Ü9-Ü10; AR Ü10-ÍÜ12; PR i 13 - i 15; VR i 8 - i 9; LLS
42-50; TS about 22; GR more than 40 long rakers, 31-33 vertebrae.
Pigmentation. The body is silvery on the dorsum and sides, but
(in life) the chest, pectoral, pelvic and anal fins are cherry red,
particularly during the breeding season. The anal and caudal fins are
sometimes edged in black.

Figure 262. Apure drainage distribution of Curimata cerasina.

CUENCA DEL RIO APURE
685

686
Distribution and Natural History
Range: It is known from the Orinoco Basin.
Apure distribution: Map: fig. 262. Curiously, it is absent from
the westernmost portion of the drainage. It occurs in the piedmont and
both high and low llanos.
Habitat. It lives in both lotic and lentic Whitewater habitats,
and is usually taken over muddy substrates.
Abundance. COMMON.
Number of specimens examined. 253 from 69 collections.
Food. DETRITIVORE. It feeds on algae, diatoms, bacteria and
protozoans from mud and detritus.
Reproduction. Probably Strategy r2. Ripe females were found from
late March to April, and juveniles (17-60 mm SL) were taken from
floating and emergent vegetation from July and August (Vari 1983).
Migration. It is large enough to accompany the coporo Prochilodus
mariae on its annual migrations, but this has not been documented.
Importance. It has some potential as an ornamental, but grow too
large for aquaria.
Curimatel1 a bolivarensis (Steindachner) 1910
Curimata - Coporito
Fig. 263. Map: fig. 264. Couplet 2a.
Generic synonymy
Curimatel1 a Eigenmann & Eigenmann 1889b:415 (type species: Curimatus
1 epidurus Eigenmann & Eigenmann 1889b, by subsequent designation of
Eigenmann 1910:420); Fowler 1950:295 (synonymy).
Apolinarel 1 a Fernández Y. 1948 (type species: Curimatus meyeri
Steindachner 1882, by original designation).

687
Figure 263. Curimatel1 a bolivarensis.

688
Walbaunina Fernández Y. 1948 (type species: Curimatus dorsalis Eigenmann
& Eigenmann 1889b, by original designation).
Lepipinna Fernández Y. 1948 (type species: Anodus alburnus Müller &
Troschel 1845, by original designation).
Specific synonymy
Curimatella bolivarensis Steindachner 1910:265 (type locality: Orinoco
River at Ciudad Bolivar), seen; Schultz 1944b:249 (key).
Curimatella bol i várense Mago L. 1970:75.
Etymology. CURIMATELLA = a diminutive of Curimata; BOLIVARENSIS =
from Ciudad Bolivar, Venezuela.
Description
Illustrations. Fig. 263; Román 1985:134.
Diagnosis. This curimatid is distinguished by having the caudal
fin almost entirely scaled, and by the presence of a large black blotch
on the caudal peduncle that is separate from the lateral stripe (if the
latter is present).
Size. It seldom exceeds 80 mm SL.
Morphology. See previous account for C_¡. argéntea.
Counts. DR ii9; AR i ii7; PR i 12; VR i8; LLS 32-36; TS 13; GR
short, more than 40 in total number.
Pigmentation. The body is silvery with a rounded black spot at
the base of the caudal fin. The fins are colorless.
Distribution and Natural History
Range. It is known from the Orinoco Basin.
Apure distribution. Map: fig. 264. This species is very scarce.
Habitat. The four points of collection for this species indicate
a wide range of habitat. One is in the piedmont over a rocky substrate

Figure 264. Apure drainage distribution of Curimatella bolivarensis.

CUENCA DEL RIO APURE
690

691
one in the high llanos, and two in blackwater low llaneran streams.
Abundance. RARE.
Number of specimens examined. 32 from 7 collections.
Food. DETRITIVORE.
Reproduction. Strategy probably r2.
Importance. It is marginally valuable as an ornamental.
Curimatella immaculata (Fernández Yépez) 1948
Curimatella - Coporito
Fig. 265. Map: fig. 266. Couplet 2b.
Specific synonymy
Lepipinna immaculata Fernández Y. 1948:27 (type locality: Obidos,
Brazil).
Curimatel1 a immaculata Géry 1977:238 (lists as synonym of Curimatel1 a
alburna); Vari 1989 (assigns to Curimatella).
Comments. This species was posthumously attributed to Eigenmann
by Fernández Yépez (1948), who found a specimen identified and named as
a new species by Eigenmann, Indiana U.M. 4316 (now presumably at CAS),
that had never been formally described. Since Eigenmann never pub¬
lished the name, authorship is attributed to Fernández Y. In the
original description Fernández Y. (1948) first spelled the specific
name as immaculata then as inmaculata in the figure legend and in later
text, but again as Lepipinna immaculata and Curimatus immaculatus in
the index. Since Fernández Yépez used the spelling "immaculatus"
first in his publication, and both Géry (1977) and Vari (1989) also
used that spelling, I have done the same here.
Etymology. IMMACULATA = without spots.

Figure 265. Curimatella immaculata.

Figure 266. Apure drainage distribution of Curimatella immaculata.

CUENCA DEL RIO APURE
(«CALA turo
en
-p*

695
Description
Illustrations. Fig. 265; Fernández Y. 1948:27.
Diagnosis. Species of the genus Curimatel1 a differ from Curimata
in having an almost completely scaled caudal fin. As its name sug¬
gests, Curimata immaculata completely lacks black markings.
Size. It grows to about 80 mm SL.
Counts. DR ii9; AR i ii8; PR i 12; VR i8; LLS about 35; TS 12; GR
more than 40 short rakers (total).
Pigmentation. There is no dark black pigmentation of any kind in
this species, though a brownish lateral stripe is often present in
preserved material.
Distribution and Natural Hi story
Range. It is known from the Orinoco and Amazon basins.
Apure distribution. Map: fig. 266. It is most common in the low
llaneran areas of northern Apure state and the Aguaro River system.
Habitat. It has been found mostly in blackwater streams, except
for two isolated sites in the Andean piedmont.
Abundance. COMMON.
Number of specimens examined. 2809 from 103 collections.
Food. DETRITIVORE.
Reproduction. Strategy probably r2.
Importance. Curimatel1 a immaculata is a peaceful fish in aquaria
and so is marginally valuable as an ornamental.
Cyphocharax soilurus (Giinther) 1864
Curimata - Coporito
Fig. 267. Map: fig. 268. Couplet 7b.

696
Figure 267. Cyphocharax sdí1urus.

697
Generic synonymy
Cyphocharax Fowler 1906 (type species: Curimatus spilurus Giinther 1864).
Xyrocharax Fowler 1913b (type species: Curimata stigmaturus Fowler
1913b, by original designation).
Hemicurimata Myers 1929 (type species: Curimata esperanzae Myers 1929,
by original designation).
Curimatoides Fowler 1940 (type species: Curimatoides ucayalensis Fowler
1940, by original designation).
Cruxentina Fernández Y. 1948 (type species: Curimata hypostoma hastata
Allen 1942 |n Eigenmann & Allen 1942, by original designation).
Specific synonymy
Curimatus soilurus Giinther 1864:288 (type locality: Essequibo, British
Guiana), seen; Steindachner 1879:5 (Ciudad Bolivar, Venez.); Eigenmann
1922:263, pi. 34, fig. 1 (British Guiana).
Curimata spilura Schultz 1944b:249 (key); Géry 1977:235.
Etymology. CYPHO = from the Latin "cifra" for zero, probably in
reference to the lack of teeth; CHARAX = as in the genus Charax one of
the first characid genera to be described, and a common root name
formation; SPIL = spot, URA = tail, referring to the black spot on the
caudal peduncle.
Description
Illustrations. Fig. 267; Géry 1977:226, 233; Román 1985:138.
Di agnosis. This fish is distinguished from the very similar C.
argéntea by the lack of a spot in the dorsal fin. It has about 33-36
lateral-line scales. The caudal fin is scaled only near its base.
Size. This is a small to medium curimatid that seldom exceeds 100
mm SL.

698
Morphology. See account for S. argéntea. Günther remarked on the
different morphologies observed in the two specimens he used to describe
this species. He stated "The two specimens on which I have founded
this species differ most remarkably in the form of the body, although
they agree in every other point, so that there can be no doubt that they
are of one species. One has the body elevated like a C. cvprinoides.
its depth being two-fifths of the total length (without caudal); the
other has the body oblong, its depth being contained thrice and a third
in the total. Both are of the same length." I have observed similar
differences in the Apure specimens, and suspect that sexual dimorphism
might explain the above phenomenon.
Counts. DR Ü9-Ü10; AR i i i8; PR i 12; VR i8; LLS 33-36; TS 12; GR
short about 35 in number (total).
Pigmentation. The body is silvery with the scales sometimes
outlined with black, a faint lateral band and a dark black dot on the
caudal peduncle. The fins are colorless.
Distribution and Natural History
Range. It has been reported from most of South America, in the
Guianas, the Orinoco, and the Amazon basins. However, this "species" is
probably a group of sister species, and so each would have a more
limited distribution.
Apure distribution. Map: fig. 268. This species is most abundant
in Barinas state, in the Suripá River system; in northern Apure state
near Mantecal, and from the Aguaro River system in the state of Guárico.
Habitat. It is usually found in streams or ponds with clear or
black waters, often amidst aquatic vegetation.
Abundance. COMMON.

Figure 268. Apure drainage distribution of Cyphocharax spilurus.

10*
9*
••
CUENCA DEL RIO APURE
CICALA litre*
700

701
Number of specimens examined. 2291 from 68 collections.
Food. DETRITIVORE.
Reproduction. Strategy probably r2.
Importance. The are marginally valuable as an ornamental.
Potamorhina altamazonica (Cope) 1878
Manamana
Fig. 269. Map: fig. 270. Couplet 4a.
Generic synonymy
Potamorhina Cope 1878:685 (type species: Curimatus (Anodus) pristigaster
Steindachner 1876, by original designation); Fowler 1950:298
(synonymy).
Gasterotomus Eigenmann 1910 (type species: Anodus 1atior Spix jn Agassiz
1829, by original designation).
Gasterostomus Fernández Y. 1948 (incorrect spelling).
Suprasinelepichthys Fernández Y. 1948 (type species: Curimatus 1aticeps
Valenciennes 1849, by original designation).
Potamorrhina Braga & Azpelicueta 1983 (unjustified modification of
Potamorhina).
Specific synonymy
Curimatus altamazonica Cope 1878:684 (type locality: Peruvian Amazon).
Anodus laticeps Eigenmann & Allen 1942:300.
Etymology. P0TAM0 = river, RHINA = nose, (significance obscure);
ALT = upper, AMAZONICA = from the Amazon.
Description
Illustrations. Fig. 269.
Diagnosis. This is the only curimatid in the Apure drainage with
such small scales, more than 90 in the lateral-line series. The scales

702
Figure 269
Potamorhina altamazonica

Figure 270.
Apure drainage distribution of Potamorhi na altamazonica.


705
are so small that they are difficult to see without magnification.
The body and fins completely lack black markings.
Size. This is a large curimatid that reaches over 150 mm SL.
Morphology. The head and body are somewhat elongate, and the fins
somewhat long for a curimatid.
Counts. DR iilO; AR i i i 12; PR i 16; VR i8; LLS about 120; TS about
47; the gill rakers are replaced by a membranous flap.
Pigmentation. This fish is plain silvery with colorless fins.
Pistribution and Natural Hi story
Range. It is known from the Orinoco and Amazon basins.
Apure distribution. Map: fig. 270. It is known only from the low
11 anos.
Habitat. This species has been collected mostly from the main
channels of larger rivers in the low llanos, over muddy substrates. It
is more riverine than many Apure drainage curimatids.
Abundance. RARE.
Number of specimens examined. 16 from 4 collections.
Food. DETRITIVORE.
Reproduction. Strategy probably r2. It is probably an egg
scatterer with no parental care, with an annual spawning bout at the
beginning of the rainy season.
Psectrogaster ci 1iatus (Müller & Troschel) 1845
Manamana
Fig. 271. Map: fig. 272. Couplet 6b.
Generic synonymy
Psectrogaster Eigenmann & Eigenmann 1889a:412 (type species:
Psectrogaster rhomboides Eigenmann & Eigenmann, by original designa¬
tion) .

706
Figure 271. Psectroaaster ciliata.

707
Pseudopsectrogaster Fernández Y. 1948 (type species: Psectrogaster
curviventris Eigenmann & Kennedy 1903, by original designation).
Hamatichthvs Fernández Y. 1948 (type species: Anodus ciliatus Muller &
Troschel 1845, by original designation).
Semelcarinata Fernández Y. 1948 (type species: Curimatus rutiloides Kner
1859, by original designation).
Specific synonymy
Anodus ciliatus Müller & Troschel 1845:26, pi. 4, fig.4. (type locality:
Essequibo River, British Guiana).
Curimatus ciliatus Castelnau 1855:58; Eigenmann 1912:268 (synonymy).
Psectrogaster ci1 i ata Eigenmann & Eigenmann 1889:413 (alternative
spel1ing).
Plectrogaster (error) ciliata Berg 1897:278.
Psectrogaster ciliatus Eigenmann 1910:420.
Curimata ciliatus Fowler 1906:305.
Curimata ciliata Fowler 1950:279; Géry 1977:228.
Curimatus esseouibensis Günther 1864:294 (Essequibo); Eigenmann &
Eigenmann 1889:23. Eigenmann & Eigenmann 1891:48.
Curimatus rutiloides Cope (not of Kner) 1872:258 (Ambyiacu).
Curimatus cyprinoides (not of Linnaeus) Cope 1872:291.
Curimata abramoides Taphorn & Lilyestrom 1984:55 (biology).
Etymology. PSECTR0 = sawlike, GASTER = belly; CILIATUS = refers to
the ctenoid scales of this species.
Description
Illustrations. Fig. 271; Géry 1977:228 (as Curimata ciliata).
Size. This is one of the larger curimatids in the Apure
drainage. It reaches over 170 mm SL.

Figure 272. Apure drainage distribution of Psectrogaster ciliatus.

CUENCA DEL RIO APURE
MtHtfjA A
709

710
Morpholoqy. This is a robust and very deep-bodied fish. The
strongly ctenoid scales are small in the area in front of the dorsal
fin on the upper sides but are progressively larger from there down to
the area in front of the pelvic fins.
Counts. DR Ü9- iilO; AR i i i 8; PR i 14; VR i8; LLS about 50-61; TS
22; the gill rakers are replaced by membranous flap.
Pigmentation. This fish is plain silvery, with colorless fins.
Distribution and Natural History
Range. It is known from the Orinoco and Amazon basins.
Apure distribution. Map: fig. 272. It is so far known only from
near the Apure modules, west of Mantecal in northern Apure state.
Habitat. It is known only from a blackwater stream in the low
11 anos.
Abundance. UNCOMMON.
Number of specimens examined. 20 from 12 collections.
Food. DETRITIVORE.
Reproduction. Probably strategy r2. It is probably an egg
scatterer with no parental care, high fecundity and annual spawning at
the beginning of the rainy season.
Steindachnerina sp.
Yellowtail Curimata - Coporito
Fig. 273. Map: fig. 274. Couplet 5a.
Generic synonymy
Steindachnerina Fowler 1906 (type species: Curimatus trachvstetus Cope
1878, by original designation).
Rivasella Fernández Y. 1948 (type species: Curimatus melaniris Fowler
1940, by original designation).

711
Figure 273. Steindachnerina sp.

712
Curimatorbis Fernández Y. 1948 (type species: Curimatus atratoensis
Eigenmann 1912b, by original designation).
Comments. I had placed this species in the genus Cruxentina
Fernández Y. (1948) but that genus is now considered (Vari 1989) a
junior synonym of Cyphocharax. But, because I have not examined its
osteology, and because it lacks the caudal spot and corporal pigmenta¬
tion attributed to members of Cyphocharax. I assign it here tentatively
to Steindachnerina. It has several other unusual features, such as a
slender, elongate body, an elevated lateral-line scale count, and a
pigmented (yellow and black) caudal fin. The other possibility is that
it is a species of Cyphocharax. Both genera were delimited by Vari
(1989). This form probably represents a new species.
Etymology. STEINDACHNERINA = after F. Steindachner, an Austri¬
an ichthyologist who pioneered research in South American fishes.
Description
Illustrations. Fig. 273.
Piagnosis. This is the only Apure drainage curimatid with a
strictly inferior mouth. It is further distinguished by having about
50 scales in the 1ateral-1ine, and the caudal-fin scaled only near its
base.
Size. It usually does not exceed 100 mm SL.
Morphology. The body is somewhat more elongated than in other
Apure curimatids. The tail is large, and deeply forked.
Counts. DR ii9; AR i ii7; PR i14; VR i8; LLS 50; TS 16; GR about
37.
Pigmentation. The body is silvery, and the dorsal fin usually
has a black dot. The caudal fin (in life) is often marked with parallel

Figure 274. Apure drainage distribution of Steindachnerina sp.

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715
gray (to black) and yellow streaks along the upper and lowermost rays.
Distribution and Natural History
Range. It is probably restricted to the Orinoco Basin.
Apure distribution. Map: fig. 274. This species is most common in
the higher portions of the llanos and in the piedmont.
Habitat. It occurs in lotic habitats from piedmont to low llanos,
in both white and black waters, and from large rivers as well as small
streams.
Abundance. COMMON, but much less so than most other curimatids.
Number of specimens examined. 182 from 27 collections.
Food. DETRITIVORE.
Reproduction. Strategy probably r2.
Importance. It is potentially valuable as an ornamental due to its
peaceful nature, colorful tail, and a disposition to clean detritus from
the bottom of its tank.
Steindachnerina argéntea (Gill) 1858
Curimata - Coporito
Fig. 275. Map: fig. 276. Couplet 7a.
Specific synonymy
Curimatus argenteus Gill 1858:62 (type locality: Trinidad), seen;
Eigenmann 1920:9 (El Concejo, Rio Tiquirito, Maracay y Río Bué,
Venez.); Pearse 1920:21 (Mouth of Río Bué, Lago Valencia in Maracay,
Venez.); Eigenmann 1922:230 (Villavicencio, Colombia; Rio Tiquirito,
near Caracas, Rio Bue near Maracay, Venez.).
Curimata argéntea Schultz 1944b:249 (key, description); Fowler 1931:407
(Pitch Lake at Guanoco, Venez.); Román 1985:138 (description, photo);
Géry 1977:235 (key).

716
Figure 275. Steindachnerina argéntea

717
Curimatus metae Eigenmann 1922:230 pi.42, fig. 1 (type locality: Que¬
brada Cramalote (probably a misspelling of Gramalote), Villavicen-
cio, Colombia).
Comments. There could be more than one species included here
since no attempt was made to distinguish local populations. The only
criteria I used to separate this species was the presence of a black
blotch in the dorsal fin, but both S. metae and S. argéntea are said to
have a dorsal spot. In Vari’s (1983) preliminary key, S. argéntea is
said to have a small caudal spot that doesn’t extend onto the caudal
peduncle, whereas S. metae is said to have that spot large and extend¬
ing onto the caudal peduncle. I have observed a wide range of varia¬
tion in the size of this spot from the same population of curimatids
and in some it even extends forward as a lateral stripe. From observa¬
tions of fish in aquaria it is evident that spot size can vary with the
"mood" of the fish. The effects of preservation are also a factor to
be considered. Either there is a complex sympatric community of curim¬
atids at most sites or (more likely) this character is simply quite
variable.
I have included S. metae as a synonym of C. argéntea because no
characters useful to distinguish the two were included by Eigenmann in
his original description. He simply stated that S^_ metae greatly
resembles S. argéntea, differing only in shape of the caudal peduncle
and coloration. However, he did not say how caudal peduncle shape
differs, and described a color pattern common to many species. If S.
argéntea is restricted to Trinidad, then the name S. metae would be
available for the Orinocan species.
Etymology. ARGENTEA = silvery.

718
Description
Illustrations. Fig. 275. Géry 1977:232 (The fish pictured as
"Curimata eleqans-qroup" is similar to this species; Román 1985:138.
Diagnosis. This curimatid can be recognized by the following
combination of characters: caudal peduncle with a spot, or horizontally
extended blotch; dorsal fin with a black dot; caudal fin scaled only at
its base; lateral-line scales about 39.
Size. This is a small species that seldom exceeds 90 mm SL.
Morphology. The body is stocky, and fairly broad for the overall
size of the fish, with a deep, robust caudal peduncle. The head and
anterior portion of body are conical and the mouth is terminal. The
scales are large, and easily visible without magnification. The pec¬
toral fins are placed low on the body and the dorsal fin in positioned
on the anterior half of the body. A well developed adipose fin is
present. The caudal fin is forked, with equal lobes.
Counts. DR iilO; AR i ii8; PR ill; VR i7; LLS about 39; TS 13; GR
a total of 38 short rakers on the outer upper and lower arches.
Measurements. GBD 48-56% SL; HL 30% (in smaller specimens) to
50% SL (larger ones); 10 44-50% HL.
Pigmentation. The body is silvery, often with a horizontally
elongated spot on the caudal peduncle. The dorsal fin has black blotch
near the middle of its base. The other fins are gray to colorless.
Distribution and Natural Hi story
Range. It occurs on the island of Trinidad and in the Orinoco
Basin.
Apure distribution. Map: fig. 276. It occurs throughout the
drainage.

Figure 276. Apure drainage distribution of Steindachnerina argenteus.


721
Habitat. It is ubiquitous, and lives in a wide variety of aquatic
habitats, but is most common in quiet waters with muddy bottoms, such as
prestamos and floodplain lagoons.
Abundance. ABUNDANT. This is the most common curimatid in the
Apure drainage.
Number of specimens examined. 4229 from 314 collections.
Food. DETRITIVORE. It feeds on algae, diatoms, bacteria and
protozoans from mud and detritus. (K. Winemiller, pers. com.).
Reproduction. Strategy: r2; fecundity: 3528 eggs/female; egg
size 0.45 mm. This species matures in about one year, and spawns
during the first two months of the rainy season. Individuals probably
spawn only once or twice during that time (Winemiller & Taphorn 1989).
Importance. It is used as bait by commercial fishermen.
Cvnodontidae
The fishes of the family Cynodontidae, called "payaras" in Vene¬
zuela, are all specialized predators adapted to life near the water’s
surface in large rivers and streams. This small family consists of
only four species in two genera (three species of which are present in
the Apure drainage). Their large, oblique, superior mouths contain
incredibly elongated lower canine teeth (located near the symphysis of
the dentaries) that characterize this family and make them easy to
recognize. Holes in the upper jaw receive these teeth to allow the
drawbridge-like mouth to close. The body plan in this group emphasizes
speed. The keeled ventrum characteristic of the payaras serves to
reduce water resistance during the rapid pursuit of prey. The low-
placed, enlarged pectoral fins provide maneuverability for rapid upward

722
attacks and braking. The eye is very large. The head and highly
distensible mouth are designed for the manipulation of piscine prey.
Their main "fishing gear" is a large pair of canines that are used to
impale the prey on the first pass. After resistance stops, fish are
disengaged from these enameled gaff hooks, flipped around and swallowed
whole, usually head first. The main canines are reinforced by numerous
additional canines in both jaws, as well as spinous gill rakers.
Payaras are common predators present in most llaneran rivers. They
readily take baited hooks and even artificial lures, especially silver
spoons, but are hard to land because of their hard bony mouths. They
are not much valued as food fish because of the numerous small inter¬
muscular bones in the flesh.
Key to the Species of Cvnodontidae in the Apure River Drainage.
la. Origin of dorsal fin well in advance of anal origin...
Hydrolvcus scomberoides (fig. 277)
lb. Origin of dorsal fin over or behind anal origin.... ...2
2a. (lb) Anal fin longer, with more than 70 rays; body short, with a
"sawed off" look, the greatest body depth about 3.5 in SL (fig.
279); distance between dorsal fins almost twice head length;
lateral-line scales about 120...
Rhaphiodon gibbus (fig. 279)
2b. Anal fin shorter, with fewer than 50 rays; body very elongate,
greatest body depth about 4.5 in SL (fig. 281); interdorsal
distance only about half head length; lateral-line scales 125-
150...
Rhaphiodon vulpinus (fig. 281)

723
Species Accounts.
Hydro!vcus scomberoides (Cuvier) 1817
River Wolf - Payara
Fig. 277. Map: fig. 278. Couplet la.
Generic synonymy
Hydro!vcus Müller & Troschel 1844:93. (type species: Hidrocvon
scomberoides Cuvier 1819, by monotypy; Muller & Troschel 1845:18;
Eigenmann 1910:444; Eigenmann 1912:396.
Cynodon (part) Steindachner 1882 (1883):15.
Specific synonymy
Hidrocvnus scomberoides Cuvier 1817:168 (type locality: rivers of
Brazil), seen.
Hydrocvon scomberoides Cuvier 1819:357, pi. 27, fig. 2 (type locality:
rivers of Brazil).
Hvdrolvcus scomberoides Müller & Troschel 1845:19; Müller & Troschel
1848:19, pi. 5, fig 2.
Cynodon scomberoides Valenciennes j_n Cuvier & Valenciennes 1849:324;
Peters 1877:472 (Calabozo, Venezuela).
Hidrolvcus scomberoides Cope 1872:292.
Hvdrolicus scomberoides Eigenmann & Allen 1942:273.
Comments. The only other species of this genus, H. pectoral is,
occurs in southern Venezuelan in Amazonas Territory. It has fewer
lateral-line scales (90 vs 100-125), and more branched anal-fin rays
(43-46 vs 33-40) than H. scomberoides.
Etymology. HYDRO = water, LYCUS = wolf; SCOMBEROIDES = like
Scomber a genus of mackerel, probably in reference to their overall
shape and silvery coloration.

724
Figure 277. Hydro!vcus scomberoides.

7 25
Description
Illustrations. Fig. 277; Géry 1977:296; Novoa et al. 1982:fig.
11; Román 1985:153.
Diagnosis. This species has the largest canine teeth of any fish
known from the Apure drainage. It can be distinguished from members of
the genus Rhaphiodon by means of fin position. The dorsal-fin origin
is well in advance of that of the anal fin in this species, but over or
behind the anal-fin origin in Rhaphiodon. See Comments above.
Size. This species reaches at least one meter in total length.
Novoa et al. (1982) reported a specimen of 560 mm TL with a weight of
about 2 kg. This low weight for such a long fish is a reflection of
the slender and elongate shape of the species.
Morphology. Although the body is elongate and compressed in
relation to fishes of most other families, this is the most robust of
the three payaras present in the Apure drainage. The mouth is hinged
like a drawbridge. The huge canines of the lower jaw slip into the
roof of the skull when the mouth is closed and the other teeth are also
conical or caniform and arranged in a single row on both jaws. The
dorsal-fin origin is anterior to the anal-fin origin, but posterior to
the pelvics. The pectoral fin is set very low on the body, its tip
reaches the pelvic-fin insertions. The anal and caudal fins are
scaled. The caudal fin is particularly large, evenly lobed, not forked
and usually heavily abraded (or perhaps nipped by piranhas?) and round¬
ed. The lateral line is complete, the scales enlarged and obvious.
The scales are ctenoid, with serrated edges.
Counts. DR 12; AR 33-40.
Measurements. HL 4.5 in SL; GBD 3.5-4.0 in SL.

726
Pigmentation. The body and head are silvery There is a large
black blotch behind the opercle. The dorsal, pectoral and ventral fins
vary, they are sometimes plain translucent white or gray, or white with
black edges, or even solid black. The adipose fin is often orange and
black.
Distribution and Natural History
Range. It is known from most of tropical South America.
Apure distribution. Map: fig. 278. It is found in most large
llaneran rivers and adjacent floodplains.
Habitat. In the Apure drainage, adults of this species have been
found mostly in or near large muddy rivers and creeks. Juveniles were
taken in the floodplain adjacent to the larger rivers and sometimes
from ponds in flooded savannas.
Abundance. COMMON.
Number of specimens examined. 80 from 14 collections.
Food. CARNIVORE. Piscivore. Goulding (1980) found fish and
scales in the stomach of this and the closely related H. pectoralis.
Loose scales probably are a result of failed attacks, since the denti¬
tion of this species would seem to preclude scale eating as a usual
feeding method.
Reproduction. Strategy: r2. In the aquatic vegetation along the
shores in the Apure River, small juveniles (30-60 mm SL) were found in
May (the beginning of the rainy season). Novoa et al. (1982) reported
the greatest abundance of mature individuals in September (late wet
season) for the middle Orinoco fisheries. Thus, breeding may occur at
different times in different populations of this species. In the
Apure drainage, breeding occurs during the late dry season, or with
the first rains in April-June.

Figure 278. Apure drainage distribution of Hydrolycus scombe raides.

Etc I 2 000000
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CO

729
Migrations. They are probable, but not documented.
Importance. It is fished commercially in the Orinoco delta, but
is not much valued because of the numerous small intermuscular spines
in the flesh, and a tendency to spoil rapidly (Novoa et al. 1982). It
is not common in the Apure drainage fish markets.
Rhaphiodon gibbus Spix 1829
Humpbacked River Wolf - Payara Chata
Fig. 279. Map: fig. 280. Couplet 2a.
Generic synonymy
Rhaphiodon Agassiz 1829:76 (type species: Rhaphiodon vulpinus Agassiz
1829, by subsequent designation of Eigenmann 1910:444).
Raphiodon (error) Müller & Troschel 1844:93.
Hydropardus Reinhardt 1849:46 (type species: Hvdropardus rapax
Reinhardt, by monotypy).
Rhaphiodontichthys Campos 1945:472 (type species: Cynodon vulpinus Spix,
by monotypy).
Cynodon Spix in Agassiz 1829:76 (type species: Rhaphiodon gibbus Agassiz
= Cynodon gibbus Spix, by subsequent designation of Eigenmann 1910:444).
Camposichthys Travassos 1946:9 (type species: Cynodon gibbus Spix 1829,
by original designation.
Soecific synonymy
Rhaphiodon gibbus Spix i_n Agassiz 1829:77 (type locality: rivers of
Brazil), not seen.
Raphiodon (error) gibbus Müller & Troschel 1844:94.
Cynodon gibbus Spix in. Agassiz 1829:78, pi. 27; Eigenmann 1912:395
(description, synonymy); Fowler 1950:330 (synonymy, distribution).
Cynodon gibbum (error) Eigenmann & Allen 1942:272.

730
Figure 279. Rhaphiodon qibbus.

731
Etymology. RHAPHI = saber, ODON = toothed; GIBBUS = hump-backed.
Description
Illustrations. Fig. 279; Géry 1972:54, fig. 5, 2.
Diagnosis. The posterior dorsal-fin origin (placed over or
slightly behind the anal-fin origin) will distinguish this species from
Hydrolvcus scomberoides. It differs from R. vulpinus in having a much
shorter body, but a longer anal fin with over 70 rays (vs fewer than
50).
Size. It can reach about 350 mm SL.
Morphology. These fish are bizarre in appearance. The rear and
anterior portions of the body are normal for the family, but the cen¬
tral portion seems to be missing, giving the impression that the tail
has been stuck onto the head. The ventral anterior profile is rounded
up towards the mouth (possibly as an adaptation to making swift verti¬
cal movements to pursue prey sighted at the surface).
Counts. DR 12; AR 73-77; PR i15; VR i7; LLS 120-163; GR 18 on
lower arch.
Measurements. HL 4.75-5 in SL; GBD 3.4-3.5 in SL; EVE equal to
snout and interorbital, 3.8 in HL.
Pigmentation. This species is mostly silvery, with a rounded
humeral spot just behind the upper margin of the opercle.
Pistribution and Natural History
Range. It is known from most of tropical South America.
Apure distribution. Map: fig. 280. It occurs throughout the
11 anos.
Habitat. Adults are usually taken from larger rivers, whereas
juveniles are taken near shore or from adjacent flooded savannas.

Fi cure 280. Apure drainage distribution of Rhaphiodon gibbus.

CO
CO

734
Abundance. UNCOMMON.
Number of specimens examined. 22 from 8 collections.
Food. CARNIVORE. Piscivore. It also includes terrestrial
insects and other creatures that fall onto the water’s surface in its
diet.
Reproduction. Strategy r2. The smallest specimens of this
species (about 90-100 mm SL) were collected in August and November,
from flooded savannas at the modules of Apure. Thus, breeding proba¬
bly occurs earlier in the rainy season.
Migrations. They are probable but not documented.
Importance. It has no food value, but is sometimes sold as a
novelty in the aquarium trade.
Rhaphiodon vulpinus Agassiz 1829
Elongate River Wolf - Payara Machete
Fig. 281. Map: fig. 282. Couplet 2b.
Specific synonymy
Rhaphiodon vulpinus Agassiz 1829:76 (type locality: rivers of
Brazil), seen; Fowler 1950: 332-333 (synonymy, distribution).
Raphiodon (error) vulpinus Müller & Troschel 1844:94.
Rhaphiodon vulpnus (error) Travassos 1946:9.
Cvnodon vulpinus 1829:77, pi. 26.
Salmo tamuco Natterer in Kner 1860:55 (no locality given).
Hvdropardus rapax Reinhardt 1849:46 (type locality: Montevideo).
Rhaphiodontichthvs vulpinus Campos 1945: 472.
Comments. The generic name Rhaphiodontichthvs Campos sometimes is
still employed for this species. The generic status of the two species
of Rhaphiodon varies from author to author. Although they are quite

735
Rhaphiodon vulpinus.
Figure 281

736
distinctive, there is a tendency to lump them regardless of their
numerous differences, perhaps because the genera contain so few species.
Etvmology. VULPINUS = fox or dog-like, referring to the canine
teeth.
Description
Illustrations. Fig. 281. Géry 1977:297; Goulding 1980:184.
Román 1985:153.
Diagnosis. The position of the dorsal-fin origin (posterior to
that of the anal fin), and the relatively short anal fin (with fewer
than 50 rays) are characteristic of this species.
Size. This is a large species, reaching at least 700 mm SL.
Morphology. The is the most elongate and compressed of the
payaras. The anal and caudal fin are covered with a dense sheath of
scales. The complete lateral line is slightly raised and emphasized,
with the canals divided into dorsal and ventral branches on each scale.
Counts. DR 12; AR about 45; PR i17; VR i7. LLS about 125-170.
Measurements. GBD 4.5 in SL; the distance between he dorsal and
adipose fins is only about half the head length.
Pigmentation. The body is silvery with a faint humeral spot
behind the upper opercle. The fins are translucent, and sometimes
edged with gray.
Distribution and Natural History
Range. It is known from most of tropical South America.
Apure distribution. Map: fig. 281. It occurs throughout the
drainage.
Abundance. COMMON.
Number of specimens examined. 66 from 12 collections.

Figure 282. Apure drainage distribution of Rhaphiodon vulpinus.

LA UNIVCASlDAO Quf SlÉMeF
UNIVERSIDAD NACIONAL EXPERIMENTAL
DE LOS LLANOS OCCIDENTALES
"EZEOUIEL ZAMORA"
VICE-RECTOR A DO EN PORTUGUESA
CUENCA OEL RIO APURE

739
Food. CARNIVORE. This primarily piscivorous species also takes
aquatic insects such as mayfly larvae (Ephemeroptera) (Goulding 1980).
Habitat. It lives in large rivers and streams.
Reproduction. Strategy: r2. Juveniles (30-80 mm SL) were found
in aquatic vegetation, along the shores of the Apure River in May.
This species probably synchronizes its reproduction with the onset of
the rains, as do most llaneran fishes.
Migrations. They are probable but are not documented.
Importance. Although it attains a large size and has a good
taste, the numerous spines limit the marketability of this species.
Ervthrinidae
Erythrinids are primitive looking fishes, with a long, wide gape,
a terete body and small rounded fins. These medium to large-sized
predators are mainly piscivorous, though other aquatic organisms are
also eaten. When hooked they are quite aggressive, and frequently
inflict nasty bites on unwary fishermen. They are readily recognized
by the absence of the adipose fin, their cylindrical bodies, large
scales (only 30-45 in the lateral-line series), short rounded fins, and
large, well-toothed mouths. They have five branchiostegal rays, 10-12
anal-fin rays, and the dorsal-fin origin is anterior to the anal-fin
origin. Weitzman (1964) discussed the osteology of this family in
comparison with the superficially similar lebiasinids.
The common guabina, Hoplias malabaricus, is similar in appearance
to the North American bowfin, and in fact, fills a similar ecological
niche. Although it can survive in almost any kind of water, it is
most commonly found in the sluggish portions of streams and in lentic

740
aquatic systems. It is usually found in fairly shallow water near the
shore, where its subdued colors and the ability to sit almost perfectly
motionless for long periods on the bottom help it go unnoticed amongst
the aquatic vegetation and leaf litter. From this position it will
ambush unsuspecting small fishes.
The family is a small one, with only three genera and about five
species (Nelson 1976). Only three of these, in two genera, inhabit the
Apure drainage. Where found, they often are one of the dominant groups
of fishes in terms of biomass. This is especially true of Hop!ias
malabaricus, which is ubiquitous in the Apure drainage.
Key to the Species of Erythrinidae in the Apure River Drainage.
la. Caudal fin usually plain, with a gray basal crescent, not spot¬
ted; dorsal rays 11-12; lateral-line scales 32-37; maxilla with¬
out canines, but small conical teeth often present...
Hoplervthrinus unitaeniatus (fig. 283)
lb. Caudal fin usually spotted and blotched; dorsal rays 14-18;
lateral-line scales 38 or more; maxilla with 2 or 3 small canines
plus a series of conical teeth... Hop!ias (2 spp.)... ...2
2a. (lb) Eye large and usually bulging, its upper edge raised above
dorsal profile of head; eye diameter longer than snout and measur¬
ing only 13-15 times in SL; body color in life dark reddish
brown...
Hoplias macrophthalmus (fig. 285)
2b. Eye smaller, its upper edge well below dorsal profile of head;
eye diameter shorter than snout length and measuring 16 to 20
times in SL; body color in life usually lighter, tan to gray with
darker black and brown blotches and spots...
Hoplias malabaricus (fig. 287)

741
Species Accounts.
Hoplervthrinus unitaeniatus (Spix) 1829
Aguadulce
Fig. 283. Map: fig. 284. Couplet la.
Generic synonymy
Hoplerythrinus Gill 1896:208 (type species: Erythrinus unitaeniatus
Spix in Agassiz 1829, by original designation).
Qphiocephalops Fowler 1906:293 (type species: Erythrinus unitaeniatus
Spix in Agassiz 1829, by original designation).
Specific synonymy
Erythrinus unitaeniatus Spix i_n Agassiz 1829:42, pi. 19, (type locality:
Río Sao Francisco, Brazil), seen; Regan 1906:382 (Venez.).
Hoplerythrinus unitaeniatus Gill 1896:208; Eigenmann 1912:418 (synonymy,
description); Fowler 1950:360 (synonymy); Géry 1977:103 (key,
description).
Erythrinus 1-taeniatus (Terror) Agassiz 1829:pi. 19.
Hoploerythrinus (error) unitaeniatus Fowler 1915:531.
Erythrinus salvus Spix i_n Agassiz 1829:41 (type locality: Río Sao
Francisco); Eigenmann & Eigenmann 1891:45.
Erythrinus gronovii Valenciennes in Cuvier & Valenciennes 1846:499, pi.
585 (type locality: Cayenne, Surinam, Brazil); Peters 1877:472
(Calabozo, Venezuela).
Erythrinus vittatus Valenciennes in Cuvier & Valenciennes 1846:499, pi.
551 (type locality: Brazil, Cayenne, Surinam).
Erythrinus kessleri Steindachner 1876:596 (type locality: Bahia,
Brazil).
Erythrinus salmoneus Cope 1878:694 (type locality: Peruvian Amazon).

Figure 283. Hoplervthrinus unitaeniatus.

743
Etymology. HOPL = is from the Greek "Hoplos" for shield or
armor, ERYTHRINUS = is a genus of the Erythrinidae; UNI = one, TAENIA-
TUS = spotted.
Description
Illustrations. Fig. 283; Géry 1977:97; Novoa et al. 1982:fig.
10; Román 1985:127.
Diagnosis. This species can be distinguished from species of the
genus Hoplias by having fewer dorsal-fin rays (11-12 vs 14-18), fewer
lateral-line scales (32-37 vs 38 or more), and in lacking canine teeth
on the maxilla.
Size. Novoa et al. (1982) reported that this species can reach
350 mm TL, and a weight of 0.68 kg, but most individuals are smaller,
between 80-140 mm SL.
Morpholoqv. This is a cigar-shaped fish with a large mouth and
an elongate gape. The posterior tip of the maxilla reaches well beyond
the anterior margin of the eye.
Counts. DR 10-12; AR 11; LLS 32-37.
Measurements. The eye diameter measures about 19 times in SL.
Eigenmann (1912) reported that the head length measures 3.3-3.5, and
the body depth 4.25 in the SL; and that the eye measures 1.5 in the
snout length, 6 in the head length, and 2.75 in the interorbital width.
Pigmentation. The characteristic spot on the rear edge of the
opercle gives this species its name. There are two or three dark lines
radiating from the eye across the preopercle and continuing onto the
opercle. Spawning males can be quite colorful. The fins and opercle
take on a reddish orange to yellow hue. The body is olive green to
bluish in life (light to dark brown in preservative, darkest dorsally),

744
with a dark lateral stripe and a series or irregular dots and vertical
bars that are darkest posteriorly. The dorsal and anal fins are spot¬
ted but the pectorals, pelvics and caudal are usually plain, although
the caudal fin has a double vertical crescent near its base.
Distribution and Natural History
Range. It occurs in most of tropical and subtropical South
America and the island of Trinidad.
Apure distribution. Map: fig. 284. It is widespread in upper
and lower llanos, in both black and Whitewater systems. It is widely
distributed throughout the Apure drainage, but absent from montane
tributaries.
Habitat. It occurs in all sorts of water bodies throughout the
llanos, but is most frequently found in bajios, esteros, prestamos and
other lentic habitats. This species is known to be a facultative air
breather (Rantin & Johansen 1984), an ability that allows it to survive
in habitats that are inhospitable for most fishes.
Abundance. COMMON.
Number of specimens examined. 120 specimens from 38 collections.
Food. CARNIVORE. Adult Hoplervthrinus unitaeniatus are mostly
piscivorous, but also take prawns and aquatic insects; subadults
feed mainly on aquatic insects. This species is much more active, and
employs a hunting strategy that is totally different from that observed
in the Hoplias species, which usually lie in wait to ambush prey. This
species actively seeks out its prey, patrolling rapidly along the edge
of aquatic vegetation in ponds and hovering in the edges of the swift
current that flows through culverts under roads. Small fishes strain¬
ing against the current are actively chased down and swallowed.

Fi qure 284.
4pure drainage distribution of Hoplerythr inu s unitaeni atus.

-p*
cr>

747
Reproduction. Strategy r2; fecundity about 6000 eggs/female; egg
diameter 1.5 mm. This species reaches sexual maturity in 12 months; it
spawns annually in April to May at the initiation of the rainy season
(Winemiller & Taphorn 1989).
Eigenmann (1912) stated that "The anal fin in the breeding males
is swollen; the pocket formed by the base of the last anal ray and the
sheath of the fin is prominent; a row of scales from the base of this
pocket to the second scale above the origin of the anal have the margin
deeply incised and covered with a roll of thick skin, which is thence
continued down to the anus." I have observed a similar secondary
sexual character in Lebiasina ervthrinoides, which is now included in
the family Lebiasinidae, but which was once lumped with erythrinids.
Novoa et al. (1982) noticed that during the spawning season in
the Orinoco Delta (April-May) all specimens collected had empty stom¬
achs. This suggests that feeding may be suspended during this time.
Migrations. It is not known to make extensive migrations, but it
does move locally to avoid being trapped during time of drought.
Importance. It is consumed locally, and sold in the commercial
fishery of the Orinoco Delta. Demand is currently low in the Apure
drainage due to the poor quality of the flesh and numerous spines.
Sometimes it is sold as a novelty item in the aquarium trade.
Hoplias macrophthalmus (Pellegrin) 1907
Aimara
Fig. 285. Map: fig. 286. Couplet 2a.
Generic synonymy
Hoplias Gill 1903:1016 (type species: Esox malabaricus Bloch, by origi¬
nal designation).

748
Figure 285. Hoplias macrophthalmus.

749
Macrodon (not Schinz 1822) Müller 1842:308 (type species: Esox
malabaricus Bloch, designated by Eigenmann & Eigenmann 1889:102).
Specific synonymy
Macrodon malabaricus macrophthalmus Pellegrin 1907:26 (type locality:
Cayenne: Mélinon, French Guiana), seen.
Hop!i as malabaricus microphthalmus [error] Eigenmann 1910:448.
Hopli as malabaricus macrophthalmus Fowler 1950:362 (synonymy).
Hoplias macrophthalmus Eigenmann 1912:413-414 (synonymy, key,
description); Mago L. 1970:73; Géry 1977:102. (key, description).
Etymology. H0PLIAS = perhaps from Hoplo, Greek for shield or
armor; MACR = large, OPTHALMUS = eye.
Description
Illustrations. Fig. 285; Géry 1977:96, 96 (misidentified as FL
malabaricus. illustration of head).
Di agnosis. This species differs from H. mal abaricus mainly in
relative eye size and pigmentation. The eye is larger, and measures
only 13-15 times in the SL (vs 16 or more in H. malabaricus). The eye
usually enters the dorsal profile. This species is darkly pigmented,
and is sometimes nearly black, whereas H. malabaricus is usually light
brown or tan in life and gray in preservative.
Size. This is the giant of the genus. It reaches over 1000 mm.
Morphology. Eigenmann (1912) characterized the genus Hoplias as
follows: Caudal rounded; no occipital process; no fontanel; no adipose;
mouth large, cheeks entirely covered by suborbitals; teeth all conical;
maxilla with a canine and numerous small teeth; premaxilla with a large
canine near symphysis and a smaller one toward the sides, and numerous
conical teeth; palatines with patches of teeth, the outer series

750
enlarged; a detached patch of teeth in front of the palatines, maxilla
extending beyond the orbit; walls of swim bladder normal; supratemporal
plate single. It has large, regularly-ordered scales and a thick
cylindrical body.
This species has the eye particularly large and bulging. The
lateral line is complete with sometimes multiple canals.
Counts. DR 15-16; AR 10; LLS 43-45; 13-14 scales above and
between lateral-line scale rows on opposite sides of body.
Measurements. GBD 4.0-4.3 in SL; HL 3.0-3.3. From Eigenmann
(1912): Eye 6.5 in head in specimen of 500 mm (probably total length),
5.0 in specimens 300-400 mm, 4.0 in specimens 50-250 mm; head 2.0 in
interorbital in the largest, 1.4 in specimens 300-400 mm; 0.6 in speci¬
mens 50-150 mm. Head 3.0-3.25 in standard length; depth 4.0-4.3.
Pigmentation. This fish has a dark reddish brown body with
irregular mottling and spotted fins.
Pistribution and Natural Hi story
Range. It occurs in the Amazon and Orinoco basins, Guyana,
Surinam and French Guiana. In Venezuela this species is usually found
in black or Clearwater rivers such as the Caroni, Caura, Cinaruco,
Ventuari and upper Amazon tributaries.
Apure distribution. Map: fig. 286. It is found only in the SE
corner of the Apure drainage.
Habitat. It is exclusively restricted to black or Clearwater
streams and rivers. This species is much less common in strictly lentic
biotopes than its congener.
Abundance. RARE. In the blackwater rivers where this species
lives, we have made little fishing effort with hook and line or with
the commercial gill nets that are necessary to catch this species.

Figure 286. Apure drainage distribution of Hoplias rnacrophthalnus.

CUENCA DEL RIO APURE
IKtL
too
en
ro

753
Number of specimens examined. 2 specimens from 2 collections.
Food. CARNIVORE. It is mainly a piscivore, but also eats other
aquatic animals. It appears to be somewhat nocturnal.
Reproduction. Probably r2, similar to H. malabaricus.
Migrations. It is not known to migrate long distances, but makes
local movements to avoid being trapped in time of drought.
Importance. It is much appreciated as a food fish in eastern
Venezuela where it is more abundant, but it is not sold commercially in
the Apure drainage.
Hoplias malabaricus (Bloch) 1794
Guabina
Fig. 287. Map: fig. 288. Couplet 2b.
Specific synonymy
Esox malabaricus Bloch 1794:149, pi. 392 (type locality: "Tranquebar" =
Tropical South America), seen.
"Tareira" Marcgrave 1648:157 (name in text).
Svnodus malabaricus Bloch & Schneider 1801:397.
Macrodon malabaricus Eigenmann & Eigenmann 1889:102 (Brazil, Surinam);
Pellegrin 1899:157 (Apure River, Venez.).
Hopl ias malabaricus Eigenmann & Kennedy 1903:508 (Brazil); Fowler
1911:433 (Pedernales area, Venez.); Eigenmann 1912:414 (synonymy,
key); Pearse 1920:21 (Lake Valencia, Maracay, Venez.); Eigenmann
1920:9 (Tuy, Bué, and Tiquirito rivers, Lake Valencia area, Venez.);
Fowler 1931:407 (Caño Guanoco, Venez.); Rohl 1942:384 (Maracaibo,
Venez.); Schultz 1944b:308 (L. Maracaibo, Venez.); Mago L. 1970:73;
Géry 1977:102 (key, description); Saavedra M. 1984:1 (biology).
Hopl ias malabaricus malabaricus Fowler 1950:362 (synonymy).

754
Figure 287. Hop!ias malabaricus.

755
Svnodus tareira Bloch & Schneider 1801:398, pi. 79 (type locality:
rivers of Brazil).
Macrodon tarei ra Valenciennes i_n Cuvier & Valenciennes 1846:508
(Brazil, Venezuela [Maracaibo]).
Svnodon palustris Bloch & Schneider 1801:398 (type locality: rivers of
Brazil).
Macrodon quay i na Valenciennes i_n Humboldt 1817:179 (type
locality: ?Guyana); Cuvier & Valenciennes 1846:527 (Lake Tacarigua,
this could be Lake Valencia, Venezuela, since that lake was once
called Tacarigua. There is also a brackish lagoon called Tacarigua on
the Venezuelan coast southeast of Higuerote. Since the name guavina
(or guabina, since in Venezuela Spanish "v" and "b" have the same
sound) is still the accepted common name here for this species it is
likely that the specimens were from Venezuela).
Ervthrinus trahira Spix i_n Agassiz 1829:44 pi. 18 (type locality:
Brazil).
Macrodon trahira Giinther 1864:281 (Guyana, Brazil); Liitken 1875:184
(Venez.); Regan 1906:382, pi. 1 (Venez.); Steindachner 1874:524
(Orinoco).
Ervthrinus macrodon Agassiz 1829:43, pi. 18. (type locality: "Lago
Almada no Estado da Bahia; Río Sao Francisco," Brazil).
Ervthrinus microcephalus Agassiz 1829:44 (type locality: Rio San
Francisco).
Ervthrinus brasil iensis Spix i_n Agassiz 1829: 45, pi. 20 (type locality:
Peruaguacu, Brazil).
Macrodon brasil iensis Miiller & Troschel 1844:82.
Macrodon auri tus Cuvier & Valenciennes 1846:519 (type locality:
Montevideo, [Uruguay]).

756
Macrodon teres Cuvier & Valenciennes 1846:521 (type locality: Maracaibo
[Venez.]).
Macrodon patana Cuvier & Valenciennes 1846:522 (type locality: Cayenne).
Macrodon ferox Gill 1858 (type locality: Trinidad).
Macrodon intermedius Giinther 1864:282 (type locality: Río Cipo)
Etymology. MALABARICUS = from Malabar, apparently there was
some error in the original locality of the specimens sent to Bloch,
since Malabar is a coastal region of SW India on the Arabian sea in
Karnataka and Kerala states. The reference to "Tranquebar" given as
the type locality in the synonymy is of unknown origin.
Description
Illustrations. Mago L. 1970:259; Géry 1977:97; Román 1985:126.
Diagnosis. This species differs from H. macrophthalmus mainly in
relative eye size and pigmentation. The eye measures 16 to 20 times in
the SL, and so is relatively smaller than in H. macrophthalmus, in
which the eye diameter measures only 13-15 times. The eye does not
usually enter into the dorsal profile. This species is usually light
brown or tan in life, and gray in preservative. H. macrophthalmus is
much darker, and sometimes nearly black.
Size. This species doesn’t grow as big as the aimara, but still
reaches a respectable 400 mm SL. It is usually taken between 100-250 mm
Counts. DR 13-15; AR 10-11; LLS 37-43.
Measurements. The following data are from Eigenmann (1912): Head
length 3-3.4 in the SL; head depth 4.3 in SL. The eye diameter measures
7 times in the head length of large specimens (200 mm SL or more); 5-6
times in specimens 100-200 mm, 4.3 - 5 times in specimens 50-100 mm. The
eye diameter fits 2.4 times in the interorbital width in specimens 300-

757
400 mm, 1.6-2 in specimens 200-300, 1.5 n specimens 100-200 mm, 1-1.3 in
specimens 50-100 mm.
Pigmentation. The fins are usually spotted. As a rule, the body
is much lighter in this species than in H. macrophthalmus, gray to tan,
with irregular blotches and spots, and the abdomen whitish. However, in
some habitats the body can be quite dark, and relative eye size must be
used to distinguish the two species.
Distribution and Natural History
Range. It occurs in Central America, and in tropical and
subtropical South America as well as on the island of Trinidad.
Apure distribution. Map: fig. 288. It is ubiquitous in the lower
Apure drainage, and are absent only from high montane streams.
Abundance. ABUNDANT. This species dominates biomass figures in
many lentic habitats in the llanos, and is one of the most common
llaneran predators (Taphorn & Lilyestrom 1984).
Number of specimens examined. 649 specimens from 252 collections.
Food. CARNIVORE. It is mostly piscivorous, but also takes other
aquatic animals such as prawns and aquatic insects (Taphorn & Lilyestrom
1984; Saavedra 1984; Lowe-McConnel 1 1975, Pearse 1920). Prey are
swallowed whole. Its hunting strategy is to sit in ambush, using
protective coloring to blend in with vegetation and debris on the bottom
in shallow water near shore.
Habitat. It has been found in almost every type of biotope present
in the drainage, with the exception of the highest mountain streams,
but is especially common in temporary or disturbed areas, and lentic
systems in general. Unlike Hoplervthrinus unitaeniatus, which has
special respiratory organs that allows it to breath atmospheric air,

Figure 288. Apure drainage distribution of Hoplias malabaricus.

en
10

760
this fish is an exclusive water-breather (Rantin & Johansen 1984). Even
so, it often inhabits hypoxic water bodies. This is possible due to a
high anaerobic capacity, and special behavioral respiratory responses
such as increased breath frequency and volume, reduction of activity at
low oxygen concentrations, and physiological adjustment of oxygen uptake
through the gills. It is tolerant of low salinity water (such as in
northern Lake Maracaibo), and pollution (it is one of the few remaining
native species in the now nearly totally polluted Lake Valencia).
Reproduction. Strategy r2. This species can reach sexual maturity
in 12 months. Spawning occurs chiefly at the beginning of the rainy
season from April to June, but can continue throughout the high water
season for about five months. The female lays between 2500-3000 eggs
(average diameter 2.0 mm) in a shallow depression in shallow water near
shore where eggs are guarded by the male. Eggs hatch in about four
days, and the yolk sac is absorbed in another ten (von Ihering et al.
1928; Azevedo and Gomes 1943).
Migrations. H. malabaricus has been reported to move overland
through wet vegetation or mud at night or during rainstorms.
Importance. It is consumed locally if nothing else is available.
Gasteropelecidae
Hatchetfishes are unique in their ability to vibrate their pec¬
toral fins to sustain "flight" when jumping from the water. They get
their common name from their greatly expanded coracoid bones that form
a characteristic, hatchet-shaped keel. The combination of a superior
mouth, large pectoral fins, and a keeled body gives hatchetfishes great
maneuverability in the water, an important ability for fishes that feed

761
at the water’s surface in competition with other species. They can
often be seen just beneath the water’s surface where they patrol for
fallen terrestrial insects.
Their "flight" serves as an escape mechanism from predators since
they take to the air and scatter when startled. I have seen them leap
from the water in aquaria trying to catch insects, but it was not possi¬
ble to see if they had actually been successful in this regard.
Weitzman (1954, 1960) discussed the osteology and anatomy of the
group, as well as the taxonomy. Géry (1977) recognized only eight
species in this family, only one of which occurs in the Apure Drainage.
Species Accounts.
Thoracocharax stellatus (Kner) 1859
Silver Hatchetfish - Pechona
Fig. 289. Map: fig. 290.
Generic synonymy
Thoracocharax Fowler 1906:452 (type species: Gasteropelecus stel1atus
Kner, by original designation).
Specific synonymy
Gasteropelecus stellatus Kner 1859:17, pi. 1, fig. 2 (type locality Rio
Cujaba), seen; Pellegrin 1899:157 (Apure River, Venez.).
Thoracocharax stellatus Eigenmann & Allen 1942:268 (Orinoco); Schultz
1944b:275 (key); Géry 1977:246.
Etymology. GASTERO = belly, PELECUS = hatchet; STELLATUS = star-
like, in reference to the shiny silver color.
Description
Illustrations. Fig. 289; Géry 1977:244; Román 1985:125.

762
Figure 289. Thoracocharax stellatus.

763
Diagnosis. The body shape of this species is unique among Apure
drainage fishes. It could possibly be confused with small Triportheus,
but species of that characid genus are much more elongated.
Size. This is a small species that seldom exceeds 40 mm SL, and
is usually found at a much smaller size.
Morphology. The body is extremely compressed, with tremendously
expanded coracoids that form a trenchant keel. The lateral line runs at
an angle down the sides toward the anal fin. The mouth is superior,
with only two or three teeth in the outer row on the premaxilla. An
adipose fin is present. The base of the anal fin is sheathed in two to
three rows of scales.
Counts. DR i i i 12; AR i ii39-i ii41; PR ill; LS 20.
Pigmentation. The body is silvery. The dorsal fin has a black
spot. The other fins are colorless.
Distribution and Natural History
Range. It is widespread from Paraguay to Venezuela.
Apure distribution. Map: fig. 290. It occurs in most of the
llanos.
Habitat. It lives in quiet creeks, often forested and shady, with
cool water, and high visibility. It is frequently observed in open
pools in schools, just beneath the surface.
Abundance. ABUNDANT.
Number of specimens examined. 2874 from 165 collections.
Food. CARNIVORE. Their diet includes terrestrial insects (espe¬
cially ants), aquatic beetles, and small seeds.
Reproduction. Strategy: r2. This species reaches maturity in 12
months. It spawns once or twice during the first two months of the

Figure 290. Apure drainage distribution of Thoracocharax stellatus.

765

766
rainy season. The 0.8 mm eggs are probably scattered and abandoned.
Fecundity is about 750 eggs/female (Winemiller & Taphorn 1989).
Actual spawning behavior has not been observed in this species,
but the similar Gasteropelecus maculatus from the Maracaibo Basin repro¬
duced in a 2000 liter aquarium. The parents were fed well, mainly with
insects attracted to a light suspended over their tank at night. The
fish scattered buoyant eggs throughout the tank and although the adults
did not provide any care, they didn’t eat their young.
Importance. This fish is a highly sought as an ornamental.
Hemiodontidae
The Hemiodontidae is a small family comprising about five genera
and some twenty-five species. Neither the correct spelling for the
name of this group nor the correct taxonomic rank (family or subfamily)
has yet been determined to all ichthyologists satisfaction. Nelson
(1984) classified it as the family Hemiodontidae and included the
subfamily Parodontinae (here treated as a family) along with the Hemio-
dontinae. Vari (1989) attributed full familial rank to this group.
These fishes can be recognized by their sleek streamlined shape
and their peculiar multicuspidate teeth (which are present only on the
upper jaw in all but one species). Their eye is covered by a heavy
adipose eyelid presumably to further reduce drag in the water. A large
powerful, forked tail provides propulsion.
Hemiodontids are usually seen in the open water currents of
medium-sized, clear or blackwater streams, often near the edges of
banks of aquatic vegetation. They are swift, streamlined, strong
swimmers that generally occupy midwater to surface niches.

767
Key to the Apure Drainage Hemiodontidae.
la. Lateral-line scales more than 110; middle of side with a large
rounded black spot at level of tips of depressed dorsal fin rays,
which in some specimens continues posteriorly as a faint lateral
stripe onto the lower caudal fin lobe...
Hemiodopsis argenteus (fig. 291)
lb. Lateral-line scales 50 or fewer; no rounded black spot on middle of
side, a dark black stripe (sometimes broken into horizontally
elongate blotches) extending from below tips of depressed dorsal
fin rays obliquely onto lower caudal fin lobe...
Hemiodopsis grácil is (fig. 293)
Species Accounts.
Hemiodopsis argenteus Pellegrin 1908
One-Spot Hemiodopsis - Hemiodopsis de Un Punto
Fig. 291. Map: fig. 292. Couplet la.
Generic synonymy
Hemiodopsis Fowler 1906:318 (type species Hemiodus microlepis Kner,
by original designation).
Specific synonymy
Hemiodopsis argenteus Pellegrin 1908:344 (type locality: Orinoco), seen;
Bohlke 1955c:7 (color pattern); Mago L. 1970:76; Géry 1977:195 (key).
Types. MNHN 87-769-770 (2 specimens mentioned in original descrip¬
tion, but no holotype designated).
Comments. Species identifications are still not reliable in this
group, although Géry’s (1977) key helps identify species groups. Nelson
(1984) considered Hemiodopsis to be a junior synonym of Hemiodus. I

768
Figure 291. Hemiodopsis arqenteus.

769
have followed the system employed by Géry (1977). Neighboring drainages
have several additional species of hemiodontids. Some additional spe¬
cies are to be expected as collecting continues.
Etymology. HEMIODOPSIS = like a Hemiodus; ARGENTEUS = silvery.
Description
Illustrations. Fig. 291; Géry 1977:204 (the photo of a specimen
identified as: "Hemiodopsis microlepis-qroup" is quite similar in the
Apure species).
Diagnosis. The completely different pigmentation pattern (a round
black dot on the side, as seen in fig. 291) and the numerous small
scales in the lateral-line series (over 110 vs less than 50) make it
easy to separate this species from H. grácil is, its congener in the
Apure drainage.
Size. It can grow to about 150 mm SL.
Morphology. The head and body are streamlined and only slightly
compressed. The gill membranes free from the isthmus. The eye is
covered with a thick adipose eyelid. The teeth are multicuspid and
numerous, with a total of about 44 in the upper jaw. An elongate axil¬
lary scale is present above the pelvic fins.
Counts. DR ii9; AR i ii9; PR i 17-i18; VR ilO; LLS 118-127; TS
about 50.
Measurements. GBD 25-28% SL.
Pigmentation. This fish is silvery blue in life, but turns brown
in alcohol. It has a large, black rounded spot on the middle of the
side below the level of the tips of the depressed dorsal fin. The spot
is about 12 scales high and 12 long, and is followed by a faint lateral
stripe that extends out onto the lower caudal fin lobe.

Figure 292. Apure drainage distributi on of Hemiodopsis argenteus.

CUENCA DEL RIO APURE
1U.
too
Cae I 2 OOOOOO
■ «CAIA tur K •
^1

772
Distribution and Natural History
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 292. So far, this fish has been
found only in northern Apure state.
Habitat. It lives in black or Clearwater streams that originate
on the floodplain, in savanna woodlands.
Abundance. RARE.
Number of specimens examined. 2 from 2 collections. It is proba¬
bly more abundant (where it occurs) than our collections indicate, since
its open water habitat is difficult to sample.
Food. OMNIVORE. Its diet probably includes primarily aquatic
vegetation and some aquatic insects.
Importance. Ornamental.
Hemiodopsis grácil is Gunther 1864
Half-Stripe Hemiodopsis - Hemiodopsis Medio-rayado
Fig. 293. Map: fig. 294. Couplet lb.
Specific synonymy
Hemiodopsis grácil is (Giinther) 1864:299 (type locality: Rio Cupai,
Brazil), seen; Géry 1977:195 (key).
Hemiodus grácil is Fowler 1950:270 (synonymy).
Etymology. GRACILIS = slender.
Description
Illustrations. Fig. 293; Bohlke 1955:5.
Di agnosis. See previous diagnosis.
Size. The largest specimens taken in this study were about 90 mm
SL, it probably grows somewhat larger.

773
Figure 293.
upmindopsis gracilis.

Figure 294. Apure drainage distribution of Hemiodopsis gracilis.

10°
9o
>0
CUENCA DEL RIO APURE
Esc l 2 OOOOOO
mui Mtrict
775

776
Morphology. This fish is a sleek study in hydrodynamics. It is
elongate, fusiform and nearly cylindrical, and not much compressed. A
large fleshy adipose eyelid covers the eye. The teeth are multicuspi-
date, and number about 20 in the upper jaw.
Counts. DR Ü9; AR ii7-ii8; PR i15-i16; VR ilO; LLS 45-48; TS 12.
Measurements. GBD 17-21% SL.
Pigmentation. H. grácilis is a plain silvery fish, with a black
stripe running down the middle of its side from a point below the tips
of the depressed dorsal-fin rays and then posteriorly across the caudal
peduncle and continuing onto the lower caudal-fin lobe. The other fins
are colorless.
Distribution and Natural History
Range. It is known from the Orinoco and Amazon basins.
Apure distribution. Map: fig. 294. It has been found only in the
Aguaro River system.
Habitat. It occurs in flowing, open water of blackwater creeks
with abundant aquatic vegetation along shore. This species is usually
associated with fairly swift currents, sandy bottoms, and gallery
forest.
Abundance. RARE.
Number of specimens examined. 31 from 2 collections. It is
probably more abundant than our collections indicate, since its open
water habitat is difficult to sample.
Food. OMNIVORE. It probably feeds mostly on aquatic vegetation,
but may include aquatic insects in its diet as well.
Importance. Ornamental.

777
Lebiasinidae
This family includes about 50 species, most of which are poorly
described (Géry 1977). Weitzman (1964) studied the osteology of the
family in detail and compared it to the Erythrinidae. There is a strik¬
ing superficial similarity between members of the genus Lebiasina and
certain erythrinids.
The best known lebiasinids are the pencil fishes, now all included
in one genus, Nannostomus (Weitzman & Cobb 1975), but so far no members
of that genus have been found in the Apure drainage, although they are
present in the nearby blackwater drainages of southern Apure state and
from the Guyana Shield, and so might be expected to occur in the black-
water habitats of the Aguaro system. Only three species of lebiasinids,
(in the genera Copel la. Pyrrhulina, and Lebiasina) are known from the
Apure drainage.
The family Lebiasinidae includes two subfamilies, the Lebiasininae
and Pyrrhulininae (Géry 1977). The Lebiasininae are fishes of medium
size (to 120 mm SL), with a fairly large mouth, a long maxilla and
mostly tricuspid teeth. They are predators that inhabit small to medium
mountain streams. The Pyrrhulininae are all miniature fishes (seldom
over 40 mm SL) with small superior mouths, short maxillas, and conical
or multicuspidate incisors. They are most common in heavily vegetated
lentic habitats of the llanos.
In most species of this family, sexual dimorphism is well de¬
veloped. Males are usually more brightly colored and have larger fins
than females. Elaborate courtship rituals, and complex breeding behav¬
iors are also exhibited.

778
Key to the Lebiasinidae of the Apure Drainage.
la. Adipose fin usually present; dorsal fin sometimes with spot at
base, but no spot in center or near tip; caudal fin with black
spot at base; a medium-sized fish reaching about 150 mm SL...
Lebiasina ervthrinoides (fig. 297)
lb. Adipose fin absent; dorsal fin usually with a dark blotch or spot
in center or near tip; caudal fin lacking spot at base; a small
fish that seldom exceeds 50 mm SL... ...2
2a. (lb) A wide, dark brown, lateral stripe present from tip of snout
to base of tail; anterior margin of maxilla sharply convex,
its dorsal tip not slipping under maxilla; gill membranes
completely united, but free from isthmus...
Copel 1 a metae (fig. 295)
2b. A narrow black lateral stripe (sometimes a series of black spots)
from tip of snout to about 7 scales behind the opercle, not
continuing to dorsal fin origin; anterior margin of maxilla a
simple concave curve as in most fishes, its dorsal tip
slipping under maxilla; gill membranes united only anteriorly...
Pvrrhulina luqubris (fig. 299)
Species Accounts.
Copel 1 a metae (Eigenmann) 1914
Copella - Voladorita
Fig. 295. Map: fig. 296. Couplet 2a.
Generic synonymy
Copel 1 a Myers 1956:12 (type species: Copeina compta Myers 1927, by
original designation).

779
Figure 297. Copel 1 a metae.

780
Specific synonymy
Copeina metae Eigenmann 1914:229 (type locality: Barrigona, Rio Meta,
Colombia, Orinoco Basin), seen; Eigenmann 1922:233, pi. 20, fig. 3.
(Barrigón, Colombia).
Copeina eigenmanni Regan (in part) 1912:393 (Bogotá [this type locality
is undoubtedly an error since there are no pyrrhulines in Bogotá).
?Copeina arnoldi Schultz 1944b:308 (Caripito, Venezuela).
Types. Holotype: CAS 60494 (IU 13251a). Paratypes: 60495(26) (IU
13251); CAS-SU 24656.
Comments. Mrs. Marilyn Weitzman (pers. com.) kindly confirmed my
identification of this species (based on specimens MCNG 1757), but even
so, the identification is still tentative pending the final results of
her revision of this group.
Etymology. C0PELLA = after Edward Drinker Cope 1840-1897, North
American zoologist and paleontologist; METAE = from the Rio Meta.
Description
Illustrations. Fig. 295; Eigenmann 1922:pi. 20, fig 3.
Diagnosis. This fish is easily recognized by the overall pigmen¬
tation pattern (fig. 295), which consists mainly of a wide dark stripe
down the side. When specimens are faded, the difference in shape of the
maxilla is useful (see key).
Size. It seldom exceeds 40 mm SL.
Morphology. The anterior edge of the maxilla is sharply curved.
The teeth on premaxilla are in a single row. The gill membranes are
completely joined to each other and form a continuous free fold over
isthmus.
Counts. DR ii8-ii9; AR i i 8, Ü9,iii9; LLS 22-24.

Figure 296. Apure drainage distribution of Copella metae.


783
Measurements. HL about 25% SL; GBD 20% SL; eye diameter about
equal to the interorbital width, but longer than snout length, and
measuring almost 3.0 times in the HL.
Pigmentation. A wide, dark brown lateral stripe extends from the
snout to the base of the caudal fin, and is separated from the dark
brown dorsum by a lighter tan stripe. The ventrum is cream to white.
The dorsal fin has a black blotch, but the other fins are not notably
pigmented.
Distribution and Natural Hi story
Range. It is known from the Orinoco Basin.
Apure distribution. Map: fig. 296. This species is restricted to
the far southeastern corner of the Apure Drainage in the Aguaro River
system.
Habitat. It occurs in blackwater streams amidst aquatic vegeta¬
tion.
Abundance. COMMON.
Number of specimens examined. 64 from 9 collections.
Food. MICROCARNIVORE. Terrestrial and aquatic insects taken
near the water’s surface comprise the principal elements of the diet,
however, algae as well as small aquatic crustaceans are also included.
Reproduction. Strategy rl. Its reproductive habits are not
known, but sexual dimorphism is pronounced, thus, some sort of court¬
ship and parental care is indicated. Males have a triply curved maxilla,
sort of "S" shaped, whereas females have the lower border of the maxilla
smoothly curved. Males are also slimmer, with longer fins.
Importance. Ornamental.

784
Lebiasina erythrinoides (Valenciennes) 1849
Creole Carp - Volador or Carpa Criolla
Fig. 297. Map: fig. 298. Couplet la.
Generic synonymy
Lebiasina Valenciennes i_n Cuvier & Valenciennes 1846:531 (type species:
Lebiasina bimaculata Valenciennes, by monotypy).
Piabucina Valenciennes j_n Cuvier & Valenciennes 1849:161 (type species:
Piabucina erythrinoides Valenciennes, by monotypy).
Specific synonymy
Piabucina erythrinoides Valenciennes jn Cuvier & Valenciennes 1849:161
(type locality: rivers of Maracaibo, coast of "Parija" [probably the
Sierra de Perijá]), seen; Giinther 1864:311 (Maracaibo); Eigenmann &
Eigenmann 1891:52 (Maracaibo); Eigenmann 1922:125 (Maracaibo); Géry
1977:126 (key); Taphorn & Lilyestrom 1980:336 (synonymy of L.
pi eurotaenia with U. erythrinoides, biology).
Pi abucina pieurotaenia Regan 1903:623 (type locality: Mérida,
Venezuela); Schultz 1944b:291, fig. 36 (key, description); Géry
1977:127 (key).
Comments. Although Eigenmann & Allen (1942), Dahl (1971), Géry
(1977), and Taphorn & Lilyestrom (1980) all suggested that the genera
Lebiasina and Piabucina are not distinct, Piabucina has not "officially"
been synonymized with Lebiasina to my knowledge, and so I here propose
such a synonymy. Characters previously suggested to be of generic
value, such as presence or absence of an adipose fin, and a cellular
anterior wall to the air bladder, have been shown to occur in species of
both nominal genera, and in any event, do not by themselves warrant
generic separation. For example, among the genera of the Characidiidae

785
Figure 297. Lebiasina erythrinoides.

786
different species of Elachocharax are characterized by the presence or
absence of an adipose fin.
Etymology. LEBIASINA = small "Lebias," apparently a common name
of some fish in Valenciennes’ time; ERYTHRINOIDES = like Erythrinus. a
genus of the family Erythrinidae that is very similar in appearance to
this species.
Description
Illustrations. Fig. 297; Taphorn & Lilyestrom 1980:336; Román
1985:129 (misidentified as L. uruyensis
Diagnosis. This species is easily recognized both by the lack of
an adipose fin, and by its larger size. Juveniles resemble species of
Pyrrhulina.
Size. Males grow to about 190 mm SL, females to about 240 mm SL.
Morphology. This fish is cigar-shaped with rounded fins. The
adipose fin is almost always present, but can be lost by injury.
Counts. DR 8-10, mode 10; AR 10-12 mode 11; PR 12-17 mode 14; VR
7-9 mode 8; LS 29-35 mode 32; CPS 11-14 mode 12; PDS 14-16 mode 15; TS
6Í - 8| mode 1\.
Measurements. Mean values as thousandths of SL from Taphorn &
Lilyestrom (1980): PDL 550; HL 255; 10 94; EYE 49.
Pigmentation. In preservative the base color is brown, tan or
yellowish. A dark lateral stripe extends from behind the opercle to
the caudal peduncle, where it fades before reaching the caudal fin.
The caudal fin has a black blotch at its base, about the size of the
eye. Above the dark lateral stripe, there is a parallel light area 1£
scales high. Above this light stripe a second dark stripe extends from
above the opercle to the region of the dorsal fin. The dorsum is

787
lighter brown. The sides below the dark lateral stripe and the ventrum
are tan. The dorsal fin has a black spot near the base of the first
rays. The caudal fin is dusky, and the anal, pelvic and pectoral fins
are whitish or translucent.
In life, the fins are orange with white edges, the body is brown¬
ish green with rows of yellow and orange spots, and the lateral stripes
and caudal spots are jet black. Schultz (1944b) provided additional
details.
Distribution and Natural Hi story
Range. It is present in the Maracaibo Basin, the Apure drainage,
and some coastal drainages of Venezuela. It is probably present in
Colombia in the eastern Magdalena Basin, and the upper Meta (Orinoco)
drainage.
Apure distribution. Map: fig. 298. It is restricted to Andean
montane and piedmont streams.
Habitat. It occurs in small to medium-sized shady streams in the
Andes mountains or piedmont that typically have clear water, fast cur¬
rent and a rocky substrate.
Abundance. COMMON.
Number of specimens examined. 409 from 34 collections.
Food. OMNIVORE. Its diet includes terrestrial arthropods, mainly
insects, some aquatic insects, flowers and small fruits (Taphorn &
Lilyestrom 1980, K. Winemiller, pers. com.) It also take crustaceans and
small fishes. It patrols just beneath the water’s surface, waiting for
prey to fall in.
Reproduction. Strategy r2. This species reaches sexual maturity
in twelve months. The sexes are dimorphic, with males being more

Figure 298. Apure drainage distribution of Lebiasi na erythrinoides.

789

790
colorful, smaller, slimmer, and with larger fins. In particular, males
have longer anal fins with longer bases. It also has a row of about
seven modified scales (each with a central elevated ridge) on the sides
just above the anal fin (the function is unknown, but they are probably
used in "contacting" the female during their courtship ritual). The 1.3
mm eggs are scattered with no parental care. Fecundity is about 2700
eggs/female. Machado A.(1974) reported on their spawning in captivity
and the development of their larvae. In nature it probably spawns
during the first one or two months of the rainy season, and may spawn
twice during the season (Winemiller & Taphorn 1989).
Migrations. They are not expected since their habitat, when
undisturbed, has water all year round.
Importance. This species is consumed by mountain people, but its
relatively small size, mostly carnivorous diet, and low population
density severely reduces its value as a commercial food fish. Culture
experiments are limited by the near absence of demand for the species
outside of its native area. It cannot compete successfully in the
market with more desirable species at this time.
Pyrrhulina luqubris Eigenmann 1922
Pyrrhulina - Voladorita
Fig. 299. Map: fig. 300. Couplet 2b.
Generic synonymy
Pyrrhulina Valenciennes i_n Cuvier & Valenciennes 1846:355 (type species:
Pyrrhulina filamentosa Valenciennes, by monotypy).
Holotaxis Cope 1870:563 (type species: Holotaxis me!anostomus Cope, by
monotypy).

791
Figure 299. Pvrrhulina luqubris. The male is shown above.

792
Specifie synonymy
Pyrrhulina lugubris Eigenmann 1922:231, pi. 21, fig. 1 (type locality:
Barrigón, Río Meta; Vil 1avicencio, Camaral, Colombia (Orinoco Basin),
seen; Fowler 1950:348 (synonymy).
?Pvrrhulina semifasciata Regan 1912:390 (type locality: Bogotá).
Pyrrhulina obermulleri Myers 1926:150 (type locality: Iquitos, Peru).
Pyrrhuli na f i 1 amentosa Fowler 1931:407 (Pitch Lake at Guanoco; Caño
Guanoco, Venez.); Schultz 1944b:307 (key); Román 1985:128.
Types. Holotype: IU 15044. Paratypes at CAS: IU 15041, 15042.
Comments. Identification of the species of Pyrrhulina is still
extremely difficult. I have not carefully compared specimens from black
vs whitewater habitats to see if more than one species is present.
Eigenmann stated in the original description that this species was
"allied to P. brevis," and Géry (1977) considered it a possible junior
synonym of that species.
Etymology. PYRRHO = is from the Greek for fire or red, LINA =
small; LUGUBRIS is from the Latin for funereal, probably alluding to
the purple body color a traditional mourning color in some cultures.
Description
Illustrations. Fig. 299; Eigenmann 1922:pi. 21, fig. 1; Géry
1977:132; Román 1985:128 (as P. filamentosa).
Diagnosis. This species is distinguished by the following combi¬
nation of characters: adipose fin absent; lateral stripe short, extend¬
ing for only a few scales behind the opercle.
Size. It seldom exceeds 60 mm SL.
Morphology. The body is slightly compressed, deeper than in
Copel la. and with elongate fins. The dorsal, anal and caudal fins have

793
a basal sheath of scales. The caudal fin is strongly forked. There are
no easily-visible lateral line pores.
Counts. DR ii8; AR i ii9; PR i12-i13; VR i7; LS 22-23; TS 6, CPS
10.
Measurements. GBD about 25% SL. The HL measures 4-4.3 times in
the SL Eigenmann (1922).
Pigmentation. The body is purple in life. A narrow black stripe
extends from the snout, passes back through the eye, and continues a
few scales behind opercle. The sides are marked with several rows of
red or orange spots. The dorsal fin has a dark black blotch and is
edged anteriorly in white. The margins of the anal, pectoral and
pelvic fins are black in males.
Distribution and Natural History
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 300. It is found throughout the
piedmont and llanos.
Habitat. It is widespread throughout the llanos and Andean pied¬
mont in lentic habitats. It is usually associated with aquatic vegeta¬
tion in quiet, clear to black waters.
Abundance. COMMON.
Number of specimens examined. 2198 from 159 collections.
Food. OMNIVORE. Their diet includes terrestrial and aquatic
insects, microcrustaceans, and algae.
Reproduction. Strategy rl. This species reaches sexual maturity
during its first year of life. It reproduces during the first three
months of the rainy season. Fecundity is relatively low at only 82
eggs per female. Eggs are about 1 mm in diameter. According to Géry

Figure 300. Apure drainage distribution of Pyrrhuli na cf lugubris.

vo
Oí

796
(1977), the closely related P. brevis spawns on top of a flat stone.
The sexual dimorphism observed (generally different color patterns and
better developed fins in the more slender males) would suggest that some
sort of courtship ritual exists. This species might be considered a K-
strategist if adult size, survivorship and longevity were greater.
Importance. This peaceful and colorful aquarium fish is highly
sought as an ornamental.
Parodontidae
The Parodontidae are small fishes adapted for a benthic life in
fast currents. They have a torpedo shaped, streamlined body, a pointed
snout, an inferior or subterminal mouth, and large, low pectoral fins.
Fishes of this family lack teeth on the middle in the front portion of
the lower jaw. Although one to three small truncate teeth can be
present on the proximal end of the dentaries (back inside the mouth),
these are not obvious. The alignment and unusual shape of the teeth in
the upper jaw is also characteristic. The teeth have long slender
stems that are tipped with a wide and highly crenulated cutting edge,
with up to 20 individual minute cusps per tooth, that is well adapted
for scraping algae and occasionally aquatic invertebrates from the
rocky substrate. The teeth project straight back (horizontally) from
the premaxillary bones, and so are aligned with the long axis of the
body, instead of projecting vertically downward (or perpendicular to
the long axis) as in most fishes.
The two genera of Parodontidae are distinguished by the number and
arrangement of the premaxillary teeth. In the dozen or so described
species of Parodon there are from eight to ten teeth set in a straight

797
line from one side of the upper jaw to the other. In the three species
of Saccodon (none of which occurs in the Apure drainage), there are
only six teeth, and they are not arranged in a straight line. The
latter genus is further characterized by well developed facial bones
that encase the head in bony armor (Géry 1977).
Only two species of Parodon are present in the Apure drainage,
and one may represent an undescribed species. I have called the latter
Parodon sp. "linea." It lives in the quieter streams of the upper and
middle llanos. This habitat is somewhat unusual since most species of
Parodontidae inhabit flowing waters over rocky substrates, usually at
higher elevations.
Key to the Apure Drainage Parodontidae.
la. Sides of larger specimens with a series of 13-15 dark vertical bars
that are not usually united by a midlateral stripe; snout elongate
and conical, its length longer than the horizontal eye diameter;
mouth set well back from tip of snout; body more slender, its
greatest depth about 27-28% SL; medium-sized fishes reaching over
100 mm SL as adults... Parodon apolinari (fig. 301)
lb. Sides usually with a dark, narrow lateral stripe that extends from
the opercle to the base of the caudal-fin rays, the line either
broken into a series of dots or absent with a single large spot on
caudal peduncle; snout more rounded, its length shorter than the
horizontal eye diameter; mouth near tip of snout; body deeper,
its greatest depth about 22-24% SL; small fishes seldom exceeding
40 mm SL... Parodon sp. "linea" (fig. 303)

798
Species Accounts.
Parodon apolinari Myers 1930
Voladorito
Fig. 301. Map: fig. 302. Couplet la.
Generic synonymy
Parodon Valenciennes i_n Cuvier & Valenciennes 1849:51 (type species:
Parodon suborbitale Valenciennes, monotypy); Fowler 1950:265
(synonymy).
Apareiodon Eigenmann 1916:71 (type species: Parodon piracicabae
Eigenmann, by original designation); Fowler 1950:263; Géry 1977:202
(considered a subgenus of Parodon).
Specific synonymy
Parodon apolinari Myers 1930:66 (type locality: Guaicaramo, Rio Guavio,
Colombia); 1942:93 (Río Guárico, El Sombrero, state of Guárico,
Venez.); Schultz 1944b:287 (key, description); Géry 1977:202
(description).
Types. The species was described from a single specimen SU 23725,
(now CAS), an 89 mm male.
Etymology. PAR = equal, ODON tooth, in allusion to the even cusps
on the teeth; APOLINARI = in honor of Hermano Apolinar Maria, Director
of the Museum at the Instituto de La Salle in Bogotá, who sent the
species to Myers.
Description
Illustrations. Fig. 301; Fowler 1943:figs.4-6. Géry 1977: 208
(?misidentified as F\ suborbitale).
Diagnosis. The snout is usually longer than the horizontal diame¬
ter of the eye. The pigmentation pattern is remarkably similar to that

799
Figure 301. Parodon apolinari.

800
of young specimens of P. suborbitale, but adults are distinctively
marked.
Size. It grows to about 150 mm SL, but is usually found smaller.
Morphology. The body is robust and not much compressed. The head
is small and conical. The dorsal-fin origin is positioned anteriorly,
much nearer the snout than the caudal-fin base. The mouth is strictly
inferior, and the snout is pointed. Each premaxilla has four finely
crenelated teeth. The maxillas each have two smaller, similar teeth
near the upper end. The dentaries have one to three truncate teeth on
the sides, but are toothless along most of their exposed length.
Counts. DR ilO; AR i7; PR i14-115; VR i7; LLS 34; TS 9; CPS 16;
PDS 11.
Measurements. HL 5.3 in SL; GBD about 23-25% SL; EYE 4 in HL, 1.7
in snout, 2 in interorbital; 10 2 in HL.
Pigmentation. In preservative the snout and dorsum are brownish,
the ventrum lighter tan. The sides are marked with 13-15 vertical bars.
The fins are plain.
Pistribution and Natural Hi story
Range. It occurs the Colombian and Venezuelan Andes in the
Orinoco Basin.
Apure distribution. Map: fig. 302. It is restricted to the Andean
piedmont and montane streams.
Habitat. It lives in Andean streams and rivers with fast currents,
rocky substrates and usually clear water.
Abundance. COMMON.
Number of specimens examined. 229 from 44 collections.
Food. HERBIVORE. It feeds mainly on algae (diatoms and other

Figure 304. Apure drainage distribution of Parodon apolinari.

CUENCA DEL RIO APURE
Ct CAL A Wl/ICi
co
o
no

803
epibenthic forms), and occasionally on aquatic insects that are presuma¬
bly found among the vegetable debris that comprises most of its stomach
contents. (Alex Flecker, pers. com.)
Reproduction. Strategy probably r2. Males have tubercles on the
snout similar to cyprinids and develop a filament on the first dorsal-
fin ray (probably a secondary sexual character present only during the
breeding season). This would seem to indicate that some sort of court¬
ship ritual is involved.
Importance. Ornamental.
Parodon sp. "1inea"
Voladorito
Fig. 303. Map: fig. 304. Couplet lb.
Comments. This species is similar in overall shape and coloration
to P. suborbitale, of the Maracaibo Basin, but doesn’t seem to grow as
large as that species. In P. suborbitale the lateral stripe in larger
specimens tends to be zigzag instead of straight. There are also dif¬
ferences in snout morphology and eye size. The Apure specimens probably
represent a new species.
Description
Illustrations. Fig. 303.
Diagnosis. The snout is shorter and more rounded in this species,
its length is less than the horizontal eye diameter. The pigmentation
pattern is similar to that of young P. apolinari, but is useful for
distinguishing adults.
Size. It grows to about 40 mm SL.
Counts. DR ii 10; AR Ü7; PR i 14; VR i7; LLS 36-37; PDS 13; TS 9
1/2.

804
Figure 303.
Parodon sp. "Tinea".

805
Measurements. GBD about 27-28% SL.
Pigmentation. Pigmentation in this species is somewhat variable
but in most specimens I have examined a dark narrow lateral stripe is
present. The stripe often extends onto the bases of the central cau¬
dal-fin rays. Sometimes the stripe is divided into a series of dots,
and in some individuals it connects a series of short vertical bars.
For some specimens the pigmentation pattern is very similar to that of
young P. apolinari. and can cause confusion with the identification of
small specimens of the latter. Small individuals of this species
sometimes will completely lack the lateral stripe, and show only a
lozenge-shaped spot on caudal peduncle, which sometimes extends out
onto the central caudal-fin rays.
Distribution and Natural Hi story
Range. It occurs in the Orinoco Basin.
Apure distribution. Map: fig. 304. This species has been col¬
lected from sites in both the high and low llanos.
Habitat. As opposed to P. apolinari, which is a strictly montane
species, this still unidintified species lives in upper llaneran habi¬
tats, and is often found in slow-moving, shaded streams.
Abundance. COMMON.
Number of specimens examined. 344 from 24 collections.
Food. OMNIVORE. It probably feeds on benthic algae and aquatic
insect larvae.
Importance. Ornamental. Since this species has less of an oxygen
requirement than P. apolinari, it has more potential as an ornamental
fish. However, neither species is easily caught, and they are seldom
seen for sale in aquarium stores.

Figure 302. Apure drainage distribution of Paro don sp "linea

00
o

808
Prochilodontidae
Mago L. (1972) revised this family and redescribed the Venezuelan
species. He recognized only three genera: Prochi 1odus (comprising
about twenty total species, two of which occur in Venezuela),
Semaprochi1odus (four total species, two in Venezuela) and Ichthvoele-
phas (two species, none in Venezuela). Géry (1977) estimated a total of
some 25 species. This family includes some of the most abundant and
valuable commercial species in South America. In the Apure drainage,
they routinely take first place in terms of tons landed (Lilyestrom
1983), and enter into the diets of numerous predatory commercial spe¬
cies, especially the large pimelodid catfishes. They are also very
important in the diet of many rural Venezuelans, and in the dry season
are the main source of protein for many families. This subsistence
fishery is difficult to assess since there are no sales records, but it
probably is as large as the commercial fishery.
Prochilodontids have adapted to life on the floodplain by exploit¬
ing two rather different food sources, mud (with the associated commu¬
nity of algae, protozoans and bacteria) and plankton. They have large,
fleshy, evertible lips covered with tiny teeth, which form a disc that
is used for rasping algae from hard substrates, and for sucking detri¬
tus from the upper layers of mud. Finely divided, elongate gill rakers
permit the consumption of rather small plankters (Mago L. 1972).
Most species make spectacular migrations en masse to move from
feeding to spawning grounds or to avoid drought conditions in the dry
season. It is during these migrations that most commercial fishing
takes place. Habitat loss and deterioration caused by dams, drainage,
sedimentation, water contamination from municipal sewers, agricultural

809
pesticides, herbicides, and fertilizers, and overfishing (in a few
areas) has taken its toll on the Apure drainage fisheries. The prochi-
lodontids as obligate migrants, are among the most severely affected
(Lilyestrom & Taphorn 1978, Barbarino D. & Taphorn 1989).
The phylogenetic position and family status of the Prochilodonti-
dae has received numerous interpretations and undergone several changes.
Roberts (1973), who studied the osteology and phylogenetic relation¬
ships, came to the conclusion that these fishes are not close relatives
of the Curimatidae, but instead are more closely related to the Anosto-
midae and Chi 1odontidae. He found that prochi 1odontids are most distinc¬
tive in characters having to do with oral and branchial dentition, the
soft anatomy of the digestive system (they have numerous minute pyloric
caecae), and the only moderately coiled gut. The reproductive male’s
ability to produce sound is also characteristic for the family. However,
this position has been challenged by Vari (1983, 1989) who reexamined the
famil ies Prochi 1 odontidae, Chilodontidae, Anostomidae and Curimatidae. His
position is that the Prochilodontidae and Curimatidae are sister groups and
taken together form the sister group to the Chilodontidae and Anostomidae.
Each family is monophyletic as demonstrated by the presence of several
synapomorphies, most of which deal with special izations of the mouth struc¬
ture and of the feeding apparatus. Géry (1977) classified the Prochilodinae
and Chilodinae as subfamil ies of the Curimatidae, and recognized Anostomidae
as a distinct family. Nelson (1984) also placed Prochilodontinae as a
subfamily of the Curimatidae.
Two different spellings for the family (or subfamily), Prochilodidae
and Prochilodontidae, have been used. Géry (1977) is of the opinion that
the former is the only name in accordance with the international rules of

810
zoological nomenclature, but Vari (1983, 1989) continues to use Prochi-
lodontidae.
Key to Apure Drainage Species of Prochilodontidae.
la. Tail and anal fin plain or with rows of small dots; lateral-line
scales 49-65 (usually 53-62); scales ctenoid...
Prochilodus mariae (fig. 305)
lb. Tail and anal fin with conspicuous black horizontal or oblique
bars; lateral-line scales 45-49 (usually 46-48); scales not
cycloid... Semaprochilodus kneri (fig. 307)
Species Accounts.
Prochilodus mariae Eigenmann 1922
Coporo, Bocachica
Fig. 305. Map: fig. 306. Couplet la.
Generic synonymy
Prochi 1odus Agassiz 1829:62 (type species: Prochi 1odus argenteus
Agassiz, = Salmo marcqravii Walbaum, by subsequent designation of
Eigenmann 1910:424).
Chi 1 omyzon Fowler 1906:309 (type species: Prochilodus steindachneri
Fowler, by original designation).
Specific synonymy
Prochi 1odus mariae Eigenmann 1922:231 , pi. 20, fig. 2 (type locality:
Colombia, Barrigón, Rio Meta), seen; Mago L. 1972:48 (family review).
Prochilodus brama (not of Valenciennes) Peters 1877:472 (Calabozo,
Venezuela).
Types. Holotype: CAS 15150. Paratypes: CAS 23942(2).

811
Figure 305. Prochilodus mariae.

812
Etymology. PRO = in front of, or projecting, CHIL = lips, ODUS =
teeth, thus "teeth projecting from lips"; MARIAE = the meaning was not
explained in the original description.
Description
Illustrations. Fig. 305; Cervigón i_n Novoa et al. 1982:fig. 12;
Lilyestrom 1983:10; Román 1985:132.
Diagnosis. This is the only Prochi 1odus species in the Apure
drainage. P. rubrotaeniatus. which is present in the lower Orinoco, is
characterized by fewer lateral-line scales (41-45 vs 49-65). Prochilo-
dontids can be differentiated from the superficially similar curimatids
by the presence of fine teeth (sometimes visible only with magnifica¬
tion) on the margins of the protrusible disc-shaped, fleshy lips. j\
mariae is easily distinguished from Semaprochilodus kneri by the absence
of black stripes on the tail and the higher lateral-line scale count.
Small specimens, or old, faded prochi 1odontid specimens often lack
pigmentation. For such material lateral-line scale counts should be
used to assure correct identification.
Size. It grows to about 350 mm SL, but is usually captured much
smaller. This species enters the commercial catch when it reaches about
200 mm.
Morphology. The body is fusiform, but slightly compressed. The
lips have fine rounded teeth along their outer margins. The abdomen is
rounded, and the postventral region is trenchant but not keeled.
Counts. DR 11-12; AR 11; LLS 49-65; TS 18-22; PDS 15-22.
Measurements. GBD 2.3-3.1 in SL.
Pigmentation. The base color is silvery to bluish gray. The
dorsum is crossed by eight or more vertical bars, which are more obvious

813
under and in front of the dorsal fin. The sides have dark, horizontal
zigzag lines (between the rows of scales) which are most conspicuous
behind the level of the dorsal fin. The abdomen is white. The dorsal
and caudal fins have spots which are sometimes arranged in vertical wavy
1ines.
Distribution and Natural History
Range. It is known from the Orinoco Basin.
Apure distribution. Map: fig. 306. The coporo is ubiquitous in
the Apure drainage, though local abundance varies seasonally because of
migrations.
Habitat. The migration of this species carries it through almost
all habitats in mountains, piedmont and llanos. It passes the wet
season on the flooded savannas in lentic water bodies (esteros and
bajios). As the dry season begins, the great majority of the population
moves upstream to the Andean foothills and mountains, where it remains
until the rains return. It travels high up into the mountains and into
surprisingly small creeks with barely any water.
Abundance. ABUNDANT.
Number of specimens examined. 346 from 112 sites. This species is
so common that it was seldom preserved. Thus, the number of sites
indicated on the distribution map do not reflect its true abundance.
Food. ILIOPHAGE. The diet includes protozoans and bacteria from
mud and detritus, epibenthic algae, and plankton.
Reproduction. Strategy r2. There is no obvious anatomical sexual
dimorphism in this species (Lilyestrom 1983), but females grow larger.
It spawns annually at the beginning of the rainy season. The fish
descend from the higher regions of the drainage to spawn in midriver

814
while moving downstream. Fecundity is very high, with 300,000--
1,000,000 eggs per female. Males form schools in the center of the
river and produce a grunting sound that can be heard on shore. Females
pass into the spawning aggregate when ready to release their eggs. The
eggs are semi-pelagic and apparently float out of the river into adja¬
cent flooded areas where development proceeds rapidly (Bayley 1973).
After hatching, the young usually proceed downstream or further out onto
the floodplain. In the Apure drainage, the esteros and bajios provide
food and cover for the growing young of the year. Evidence from the
studies of Novoa et al. (1982) shows a high correlation between the
degree of flooding and the success of the year class. Evidently, the
more it floods, the more suitable habitat is available, and consequently
more juveniles survive to enter the fishery. Efforts to drain wetlands
or prevent their flooding will therefore directly reduce the number of
coporo that enter the fishery each year.
Migrations. Spectacular annual upstream migrations of thousands
of fishes occur at the beginning of the dry season. Fish travel up¬
stream into the piedmont and mountains, seeking stable water levels,
security from predators and a food source in the epibenthic algae that
grow on hard substrates when reduced flows of the dry season increase
light penetration. The coporos remain in the highlands, feeding until
the rains return. During the first month of the rains, they descend,
spawning along the way. The wet season is passed in aquatic meadows of
the floodplains where they fatten for the next cycle.
Importance. This is the most important (by weight) commercial
fish in the Apure Drainage, as well as in the Orinoco Delta (anonymous
1983).

Figure 306. Apure drainage distribution of Prochi lodus mariae.

816

817
Semaprochilodus kneri (Pellegrin) 1909
Bocachico
Fig. 307. Map: fig. 308. Couplet lb.
Generic synonymy
Semaprochi 1odus Fowler 1941:171 (type species: Semaprochi 1odus
SQuamilentus Fowler 1941 by original designation).
Specific synonymy
Prochilodus kneri Pellegrin 1909:155 (original description, type
locality: Rio Orinoco, Venezuela), seen; Schultz 1944b:265 (cited with
reservations in synonymy of S_^ laticeps.
Semaprochilodus kneri Mago L. 1972:58 (redescription, key,); Géry 1977:215
(key); Novoa et al. 1982:277; Román 1985:130.
Etvmology. SEMA = like, PROCHILODUS = the genus Prochi 1odus ;
KNERI = after Rudolf Kner, ichthyologist and naturalist.
Description
Illustrations. Fig. 307; Mago L. 1972:fig. 9; Cervigón jn Novoa
et al. 1982:fig. 13; Román 1985:130.
Diagnosis. See previous diagnosis to distinguish from P. mariae.
This fish can be confused with the zapuara (also spelled sapuara),
Semaprochi1odus laticeps. but the latter does not occur in the Apure
drainage, and usually has the opercular membrane edged in jet-black, the
entire ventral fin red to orange, and has more bands (up to seven per
lobe) in the caudal fin. It can also be distinguished by the relative
position of the pelvic-fin insertion, which is well behind the dorsal-
fin origin in S. kneri, but almost directly under the dorsal-fin origin
in S. 1aticeps.

818
Figure 307. Semaprochilodus kneri.

819
Size. It can grow to about 350 mm SL. Novoa et al. (1982) re¬
ported a specimen that measured 315 mm total length and that weighed 0.5
kg, The largest specimen recorded in their commercial records was 433 mm
TL. Most specimens are much smaller young of the year.
Morphology. The body is somewhat compressed. The dorsal profile
is straight to slightly concave. The insertions of the pelvic fins are
behind the dorsal-fin origin.
Counts. DR 11-13; AR 10-12; PR i 13-i16; LLS 45-49; scales above
lateral line 8-9; between anal and lateral line 6-7 1/2; between pelvics
and lateral line 7-9; PDS 11-13; CPS 16-20.
Measurements. GBD 2.3-2.8 in SL. 10 2-2.5 in HL.
Pigmentation. The anal and caudal fin are boldly marked with
alternating black and yellow bands (a central band and 3-4 more black
bands on each lobe of caudal are present) but the dorsal and pectoral
fins are transparent and the tips of the ventral fins may be orange, red
or yellow. The body and the edge of the opercle are usually silver, and
the opercular membrane is only weakly pigmented, not jet-black as in S.
laticeps. Small or poorly preserved specimens may appear plain silvery,
without any characteristic markings.
Distribution and Natural History
Range. It is known from the Orinoco Basin.
Apure distribution. Map; fig. 308. It is known from the Aguaro
River system of the southeastern Apure drainage.
Habitat. It occurs in clear and blackwater streams of sandy-soiled
savannas. Abundant aquatic vegetation is usually present.
Abundance. RARE. It is more common in the lower Orinoco.
Number of specimens examined. 6 from 4 sites.

Figure 308. Apure drainage distribution of Semap no ch il odus kneri.

CUENCA DEL RIO APURE
MCAIA «AAMC*
CO
ro

822
Food. ILIOPHAGE. Its diet probably includes bacteria and proto¬
zoans found in mud and detritus, and epibenthic algae.
Reproduction. Probably r2. It is an annually spawning species
that initiates reproduction with the first rains. Fecundity is very
high, as in most members of the family.
Migrations. In the Orinoco delta this species migrates along with
the zapuara from July to September (Novoa et al. 1982). The Apure
drainage populations have not been studied. I suspect that it moves in
and out of the flooded savannas of Guárico state seasonally, and then
descends the Orinoco to join with other populations further downstream.
Importance. Ornamental. It is sometimes sold as a commercial
species, but because of low abundance it is not particularly important
in the Apure drainage.

DISCUSSION
Ecological Considerations
The bulk of the autecological information gleaned from the field
data sheets is presented in the individual species accounts. Here I make
some generalizations about characiform’s adaptations to the llaneran
environment and to each other.
Physical parameters of the Apure River drainage
The Apure River, which drains 167,000 km^ and delivers an annual
average of 2,000 m^/s of water to the Orinoco, is one of Venezuela’s
largest. Its drainage is geographically widespread and includes all the
territory in the Venezuelan states of Portuguesa, Cojedes and Barinas, a
large portion of Apure and Guárico, and parts of Táchira, Mérida, Tru¬
jillo, Lara, Yaracuy, Carabobo and Aragua. This vast region includes a
wide range of aquatic habitats, and characiforms are found in most of
them. Most of the Apure’s tributaries are born in the Andes, where the
highest areas (between about 3500 and 5007 m), called páramos, consist of
cold, rocky meadows of plants specialized for life above the treeline.
The highest páramo streams are cold, usually around 5°C and are usually
devoid of native fishes, although trout have been introduced into some,
and astroblepid and trichomycterid catfishes can occur at these
altitudes. Below this cold zone most of the mountains are (or were
originally) forested. Andean gradients are steep, and the rivers that
drain them descend with a tumultuous roar during the wet season.
823

824
Few characiforms have been able to adapt to the low temperatures,
rapid currents and continuous scouring action of the substrate, but there
are notable exceptions. The saltador, Salminus hilarii, is a salmon-like
piscivore that inhabits this region, the palambra, Brvcon whitei, is
another montane species that reaches a large size (up to 4.0 kg). Both
are considered excellent sport fishes, but habitat destruction has made
them scarce. There are also a few smaller characids that have adapted to
life in the mountains, such as Hemibrvcon metae, Astvanax metae, and
Creagrutus cf beni. Although the characidiids are typically found in
riffles, only two species are found exclusively in the higher montane
regions, Characidium chupa and C. voladorita. The piedmont (roughly the
area between 100-500 m) is a fairly narrow transitional area between the
mountains and the plains. Heavily eroded hills and terraces characterize
this region, rivers slow their pace slightly as they pass through, and in
general aquatic habitats are more hospitable and harbor a wider variety
of fishes. Besides the full-time inhabitants, the mountains offer refuge
to many species during the dry season, when water velocities slow, the
rivers drop in level and decant much of their sediment load. The most
important of the part-time occupants is the coporo, Prochilodus mariae,
but many other fishes also ascend. The morocoto, Piaractus brachypomus.
tusos Schizodon isoonathus. several species of mije, Leporinus spp, all
take advantage of the dry season refuge found in the Andes.
By all accounts, the llanos dominate the drainage. They are often
divided into high and low llanos, but this distinction is somewhat
subjective. High llanos are usually defined as those areas adjacent to the
piedmont, between about 70-100 m above sea level. They are better drained,
more suitable for pastures and crops, and less susceptible to long-term

825
flooding than the low llanos (between about 45-70 m). It is in the llanos
that fish diversity and biomass reach their peaks (Mago L. 1978, Taphorn &
Lilyestrom 1984, Machado A. 1987).
The llanos are extensive, exceedingly flat alluvial plains (where a
gradient of only two to three centimeters per kilometer isn’t unusual).
Although Portuguesa and small parts of other states have soils well-
suited for agricultural crops, much of the Apure drainage is covered with
poor gravel-laden soils, and reddish, heavy clays interspersed with areas
of white sands (the latter most notably present in northern Apure state,
central and western Barinas state near Ticoporo and El Nula, and the
Aguaro system in Guárico state). When undisturbed, the soils supported a
natural terrestrial vegetation of open, dry tropical deciduous forest in
the "high" llanos and grassland savannas on the poorer, fully leached,
sandier soils. But cattle ranching, the most extensive land use in the
Apure drainage, and agriculture have caused widespread deforestation and
annual burning. Portuguesa state, which was over 90% forest before man
arrived, is only about 10% forested today.
Some climatic statistics would seem to indicate environmental
stability for the llanos, but this is an illusion. For example, the
average annual air temperature never varies much from between 26-27° C.
Daily air temperatures vary much more widely. Water temperatures vary
from lows around 23-24° in the shady forested streams near the mountains
to over 37° C. in open, shallow, flooded savannas. In reality, however,
this region suffers climatological extremes of astonishing proportions.
Characiforms, as well as most other Apure drainage fishes, are inex¬
tricably linked to the annual rainfall pulse that reverberates throughout
the aquatic and terrestrial communities. Water regulates and controls

826
their life cycles. It sets the time of their birth, and for most, the
time of their death. This annual heartbeat drives the evolutionary
forces, and casts the dice for the lottery of natural, catastrophic
selection that has shaped this remarkable assemblage of plants and
animals into wonderfully complex communities that we are just beginning
to understand.
Although annual rainfall in the llanos is, on the average, high by
North American standards, 1200 mm (47 in) to 1800 mm (71 in) per year,
the effect of these rains is even more pronounced since they are concen¬
trated in the seven or eight month wet-season from April to November.
Rainfall in some mountain areas is even higher (over three meters).
Rivers like the Caparo, Santo Domingo, Boconó, Guanare and Portuguesa
annually swell with sediment-laden torrents that overflow their banks and
flood the surrounding lowlands. In contrast, the deserts near
Barquisimeto, receive infrequent downpours that provide only 400 mm (16
in) of rain per year. Average runoff from the Apure River drainage
during a 21 month study period was measured at 361 mm/year by Saunders &
Lewis (1988a). The wide range in gradient (over 5000 m to about 43
meters above sea-level) creates an equally wide range in substrate type.
Boulders and cobbles pave Andean streams. These gradually shift to finer
gravel, sand, mud and silt as rivers leave the mountains and drop their
sediment loads on the plains. The heavy load of sediments, and abrupt
shift from very high to very low gradient causes a peculiar topography to
develop in the high llanos. The rivers flow perched atop levees of their
own making. As the waters descend from the Andes and enter the llanos,
they lose velocity and carrying capacity and drop much of their sediment
load. Through the ages these deposits form natural levees. The elevated

827
river bed thus formed is not very high, but since the natural slope is so
gentle, once they top their banks, river waters must flow many kilometers
over the plains before they can return to a channel. This phenomenon
occurred a few years ago in the town of Arismendi, which, when the
Guanare River flowed through it, was a fishing village. But the Guanare
jumped its banks during a high flood, cut a new course south of town, and
left the fishermen’s dugout canoes in Arismendi high and dry. Engineers
tried in vain to coax the roving river back into its old channel without
much success. Because the rivers wander about so, most of the llanos
form part of the floodplain at one time or another, and many abandoned
meanders remain as lakes or esteros (Ramia & Morales 1978) sometimes
interconnected by seasonal anastomosing waterways that fill with rain or
floodwaters each year in the rainy season. When the Apure River nears
the Orinoco in the area to the southeast of San Fernando de Apure, it
forms an internal delta of some 5000 km^ (Saunders & Lewis 1988a) because
of the damming effect exerted by the waters of the Orinoco. During the
peak rainy season from June to September the entire area floods and forms
a vast inland sea of slowly moving fresh water.
A visitor to this area who arrives for the first time in March
would scarcely believe that the llanos flood so extensively, for the dry
season is truly devastating, especially to the fish fauna. As we will
see, this annual climatic catastrophe plays a key role in determining
fish community structure and the life history strategies of llaneran
fishes. From December through March, almost no rain falls. The inland
sea evaporates away, grasslands parch and then burn, the earth cracks
under the searing heat, and violent winds lift ashes and dust into a hazy
sky. Sunsets usher darkness in with earthy red hues caused by suspended

828
particulate matter. Stranded fish die by the millions, providing a rich
bonanza to the terrestrial communities and especially to the wading
birds. Water is limited to main river channels, a few of the larger
creeks, lagoons and man-made reservoirs and ponds. Life concentrates
round these oases. A very narrow bottle-neck relentlessly culls fish
populations to a tiny fraction of their numbers and biomass at peak
flood. Thus, the extreme variability of climate, altitudinal gradient,
and the tendency of most llaneran rivers to go "a-roving," has created a
rich mosaic of aquatic habitats that has undoubtedly contributed to the
rich species diversity encountered here.
To better appreciate these conditions and their effects on fish
communities, one could imagine that the llanos as a part-time desert and
a part-time marsh. It is both of these extremes and passes through all
of the intermediate states each year. Classic limnological separations
between lentic (standing) and lotic (moving) aquatic habitats are diffi¬
cult to apply here because microhabitats are continually shifting. As a
result, as Roberts (1972) noted, faunal composition changes continually.
In a sampling program carried out in Caño Maporal (Matheus 1984), we
fished monthly at precisely the same site, for over a year in a tributary
to the blackwater Caño Caicara in northern Apure state. Seldom did the
monthly samples exceed forty species, and yet total number of species at
the end of the project approached 120. As conditions shifted with the
advancing seasons, different species occupied the area. Thus, with a
constant state of flux in environmental parameters and fish assemblages,
meaningful aquatic habitat classification, so necessary for environmental
planning and effective fisheries management proves frustratingly elusive.
But progress is being made.

829
Limnological investigations in the Amazon basin, pioneered by H.
Sioli (1984), have shown that there are three fundamentally different
types of water in the neotropics: black, clear and white. At first this
classification was somewhat arbitrary and subjective, based mostly on
apparent color. Blackwater streams, (better described as brown, tea-
colored, or even reddish), are stained with organic acids leached from
decaying vegetation, have almost no nutrients or other dissolved solids,
very low pH, and low productivity. Whitewaters (so called because they
are muddy and appear whitish from a distance) carry heavy sediment loads,
are usually neutral to slightly acidic or alkaline, have low visibility,
but are relatively productive. The Clearwater category is probably the
least satisfactory of the three for it can include variable stream types
such as Whitewaters that have dropped their sediments, very slightly
stained blackwaters, as well as true clear water (for example, spring
flows). While the classification does tend to oversimplify reality, it
has been a useful model to promote investigation of tropical limnology.
In the Apure drainage, the soils and waters are seldom as poor in
nutrients as those described for most of the Amazon Basin (Fittkau et al.
1975). Even though the pH of the water, hardness and nutrient content
vary with the origin, soils and vegetation of the stream system, most can
be classed into one of the three general types described above.
Whitewater streams originate in the Andes, are usually muddy, have a
slightly acidic to slightly alkaline pH, and are relatively high in
nutrients. Blackwater streams originate in the flat, sandy parts of the
lowlands such as the Guaritico system in northern Apure state, are
usually stained reddish-brown, have very low turbidity, are acidic (pH 5-
6) but not as extreme as Amazonian waters, and have low amounts of

830
dissolved minerals. Permanent Clearwater streams are not common in the
Apure drainage, but a few exist where springflows originate, and
temporarily clear streams form in the dry season when mountain streams
from forested areas decant most of their sediment load due to decreased
flow. In an area where extensive flooding is the rule, these three basic
types can be expected to mix, and they do so extensively. Some streams
in the lower portions may seasonally change from one water type to the
other as water sources vary. As a result, many llaneran fishes have
adapted to exploit a wide range of constantly shifting conditions in the
numerous microhabitats available.
More precise data have refined our understanding of tropical riv¬
ers. Since the Apure River has a relatively high sediment load (median,
233 mg/1) and a conductance between 88 and 225 uS/cm (Saunders & Lewis
1988b), it is classified as a Whitewater river. Those same authors
characterized some features of the Apure River’s water chemistry and
found discharge weighted mean concentrations for total phosphorus of 188
ug/1 , 957 ug/1 for total nitrogen, and 9.8 mg/1 for organic carbon.
Annual transport was 0.68 kg.ha'* for total P, 3.45 kg.ha'* for total N,
and 35.4 kg.ha"^ for organic C. Particulate matter accounted for 68% of
P, 54% of N, and 37% of C transport. The molar C:N:P ratio for the Apure
is 135:11:1; for the Caura (which has a much higher average runoff of
2423 mm) it is 772:47:1. Dissolved phosphorus, nitrogen and organic
carbon, were all found to rise with increasing discharge in the Apure,
but fell during the prolonged inundation phase. As the water receded,
concentrations of particulate fractions and some dissolved inorganic
fractions rose again. The pattern in the Caura was different, in part
due to the very different configurations of the drainages. The Caura

831
drains the rocky Guyana Shield, is mostly confined to its main channel
and has a very narrow floodplain. The Apure drains the geologically
young Andes, and is in contact via flooding with a very wide extensive
floodplain for most of the year. These physical variations in nutrient
and light availability lead to corresponding biotic variations.
Productivity of the Apure River.
Lewis (1988) measured the annual mean gross primary productivity
(as carbon) of the Caroni, Caura and Apure rivers in Venezuela. The
Whitewater Apure registered the highest value, 26 mg.m'^.d'l, the Caura
13 mg.nT^.d'l, and the Caroni only 4 mg.m'^.d'^. The main channel of the
Orinoco River varied from 19-43 mg.m'^.d'^. This was true in spite of
the suppression of potential production by dissolved and suspended
substances, which was calculated as 85% for the Apure, but only 40% for
the Caura and Caroni. Autotroph biomass (phytoplankton) values were also
highest in the Whitewater Apure river. Zooplankton abundance and
transport were studied by Saunders and Lewis (1988b) in the Apure River
for a fifteen month period. They found more than 50 species of
zooplankton, but nine rotifer species accounted for more than 90% of the
total density (mean, 138 individuals/1). Crustaceans comprised 46% of
the annual mean zooplankton biomass (1.9 ug C.W), but only 2% of
zooplankton numbers. Zooplankton abundance showed a marked, inverse
relationship to the river’ s water discharge rate. These values are much
higher than those reported for other neotropical rivers. For example,
Vásquez (1984) found maximum densities of rotifers of only 5.2.L*1 in the
blackwater Caroni, and Saunders & Lewis (1987) found up to 36.L"* in the
Caura.

832
Characiform adaptations to their environment.
Fishes in the Apure drainage have responded to the fluctuating
seasonal extremes of drought and flood in many different ways. Species
that live most of their lives in large river channels (eg. the
cynodontids), and those limited to high mountain streams in forested
areas (eg. small tetras like Corynopoma riisei). experience less
environmental fluctuation and have adopted appropriate life history
strategies (Winemiller & Taphorn 1989).
Perhaps the most spectacular behavioral adaptation to the seasonal
wetlands is the "ribazón," or upstream migration of the coporo, Prochi -
lodus mariae (see species account for details), with its associates such
as the palometa, Mvlossoma duriventris, the mije, Leporinus friderici,
and the palambra, Brvcon whitei.
Some species have "chosen" to physiologically adapt to the harsh
dry season conditions instead of migrating away from them. During the
drought, the water in the remaining pools is often fouled by decaying
vegetation, rotting fish carcasses of less tolerant species, and feces
from the hundreds of piscivorous birds that gather to reap the harvest
that waits, gasping for breath, near the surface in the shallows. Most
fishes try to avoid being trapped under such circumstances and move
locally (the lateral migrations of Welcomme (1979)) from drying savannas
into streams and rivers. Those caught tardy will often go to considera¬
ble lengths to escape. The guabina, Hop!ias malabaricus. is known to
travel overland in search of better pools when conditions get tough. Air
breathing, allows the agua-dulce, Hoplervthrinus unitaeniatus, to toler¬
ate conditions that are lethal to most fishes. But even these
adaptations are not foolproof, the environment is just too unpredictable.

833
The above are just a few of the classic, frequently cited (Lowe-
McConnell 1975, Welcomme 1979, Machado A. 1987, Taphorn 1988) examples of
adaptations that fishes have made to survive on tropical floodplains and
savannas. But they are the special cases, and account for very few of
the 137 characiforms that are found in the Apure drainage. What are the
rest doing? Simply stated, they play the numbers game. That is, they
reproduce by the millions, filling every available nook, niche and cranny
with fish during their heyday in the rainy season. They pay the price
when the floodplain dries up again and most of the individuals (probably
more than 99%) die. Roberts (1972) suggested that chance plays a major
role in the local distribution of fishes in the tropics. Chance is also
important in the determination of which individuals will make it through
the bottle-neck to survive. The characiforms literally place eggs in as
many baskets as possible and "hope" that somewhere among all the millions
that are born each spring, somebody will make it. Of course, many
species hedge their bets with physiological and behavioral adaptations to
adverse conditions, to get a slight edge just in case they make it
through the first crunch. But even so, it is still mostly up to chance
to determine which specific individuals will make it. Different local
conditions make year to year "predictions" of safe dry-season harbors
nearly impossible, especially for the individual fish. But the species
survive and indeed flourish because they have individuals everywhere
possible, and play the odds. As we will see, there are different life
history strategy options available to fill as many nooks and crannies as
possible with offspring.
The different life history strategies adopted by characiforms,
represent compromises between the opposing ecological pressures (for

834
example high juvenile survival vs high fecundity) that pull a species in
different directions. Winemiller & Taphorn (1989) grouped the different
types of compromises into three basic categories, rl, r2 and K strate¬
gies. Each strategy can be considered the endpoint or extreme of one
branch of a three dimensional continuum of responses, with each species’
"best answer" being potentially different from all others. This classi¬
fication is used here to denote the three tendencies among the greatly
varying reproductive strategies observed in Apure drainage fishes that we
have recognized.
The first tactic, termed the K-strategy, is characterized by a
relatively high juvenile and adult survivorship, a relatively long
generation time and long life. Fecundity in species that have adopted
this strategy is low per reproductive bout, but reproduction may be
repeated (iteroparity) as long as conditions remain adequate and food is
available. Some of these species even continue to spawn during the dry
season. On the average, these species suffer much smaller population
density fluctuations between the wet and dry seasons. K-strategists tend
to be medium to large-sized fishes. There are only a few species that
have adopted this tactic in the Apure drainage, and none of them are
characiforms. Although the red-bellied piranha, Pygocentrus caribe,
probably builds nests and guards its eggs, it has a relatively high
fecundity, and spawns but once per year as do most r2 type fishes.
I have likened the characiform’s strategy for survival to a lot¬
tery, but there is a twist. It is more like buying all of the tickets in
a lottery to be sure you win (ie, you get a few winners, but millions of
losers). Since the prize (goal) here is species survival, not money,
this strategy is very successful. All the Apure drainage characiforms

835
play it to some extent. The second, or rl-strategy, is applied to those
species with low juvenile and adult survivorship, low fecundity per
reproductive bout, but many bouts per season, a short generation time,
and population densities that fluctuate greatly between the wet and dry
seasons. About half of the Apure drainage characiforms have adopted this
strategy as the best option to an unpredictable (in the short term),
fluctuating environment with a catastrophic reduction in habitat and food
availability in the dry season (and the resulting annual "crash" in their
populations). The common tetras (called "sardinas" in Venezuela) such as
Astvanax. Moenkhausia. Hemigrammus etc. are all rl strategists. They
make up for their lack of longevity, size, and protection of their young
with high numbers of small individuals that reproduce repeatedly for as
long as conditions are favorable.
The third tactic, the r2-strategy, is characterized by very high
fecundity in an annual (one-time) spawning bout usually undertaken in
schools or groups of fishes instead of individual pairs, low juvenile
survivorship, but high adult survivorship and longevity, and large
fluctuations in population density between the wet and dry season. A
typical example of this type of response is the coporo, Prochi 1odus
mariae, the most important species of the commercial fishery. The coporo
spawns in large schools in the center of the river at the beginning of
the rainy season. Each female releases hundreds of thousands of semi-
buoyant eggs which are fertilized "en masse" by large numbers of
attendant males. The eggs are abandoned to drift out into the fertile
waters of the floodplain.
There are many concomitant adaptations to this approach to survival
of the lucky. Many fishes are particularly tolerant to low oxygen

836
levels, even if they can’t breathe air. Winemiller (1989) has shown that
many characiforms grow protuberances on their lips to allow them to
breathe the better oxygenated surface layer of water. Most species can
tolerate high temperatures that would quickly prove fatal to most tem¬
perate fishes. Many species fast for the dry season months. This is
true not only of the large predators that lay down heavy fat deposits by
consuming lottery losers, but also of smaller fishes. These adaptations
assure that the sweepstakes winners will be able to make it through the
adverse conditions that prevail.
Ichthyologists tend to focus their attention on the dry season
adaptations because it is at this time that natural selection and chance
act most devastatingly on the fish populations. At first examination,
the massive mortalities seem to bear grim tribute to the now somewhat
dated "survival of the fittest" theme. Indeed the widespread deaths at
this time calls to mind an evolutionary day of reckoning. But these are
catastrophic deaths, and although they are predictable from year to year
on an evolutionary time scale, at the individual level (the only level
where selection can act so far as anyone has been able to show), chance
plays more of a role in determining "who" makes it. We carefully study
and document the extreme adaptations that allow a few unusual species or
individuals to make it through the dry season, and while these are
extremely interesting, and many quite ingenious and amazing, these
examples tend to make us overlook the simple fact that most fish, even
those with the elaborate adaptations, simply die. They rely on the gonad
power of the lucky to replenish their numbers, and thus insure the
survival of the species, for one more year.

837
In Table 3 we can see that characiform species are about evenly
split between the rl and r2 strategies, and that none are K-strategists.
Since characiforms occupy all available habitats from mountain to llanos,
lentic and lotic, forest, savanna etc., and take advantage of all the
varied food resources available with a wide array of morphological,
physiological and behavioral adaptations, we might ask "Why are there no
large characiforms that take care of their young?", "Why are there no
characiform K-strategists?". If fishes could speak they might respond
"It’s simply not done!". In other words, phylogenetic constraints seem
to be involved. Apparently Apure drainage characiforms simply do not
have the K-strategy program included in their genetic material. We can
observe the other extreme in the family Cichlidae, almost all of which,
large or small, are K-strategists. Breaking out of these phylogenetic
constraints would seem to be much more difficult (none of the 138 species
of Characiforms in the Apure drainage have done so) than switching
habitats (eg. Prochilodus mariae and its migrations) switching diets (eg.
juvenile Pygocentrus caribe eat plankton, then switch to aquatic insects
and later small fishes) or even changing their physical morphology (eg.
species that temporarily develop lip protuberances to breath the oxygen-
rich water at the surface). Another reason that r-strategists dominate
in the Apure drainage has to do with the strongly seasonal nature of the
llanos and the annual catastrophic population crash it causes. Pianka
(1974) pointed out that K-strategists should be favored in "saturated"
environments, those with all the niches filled with species. The annual
fluctuation in most llaneran aquatic habitats probably prevents fish
communities from ever reaching equilibrium. Thus, a "poly-unsaturated"
environment is recreated each year when the rains return. R-strategists

838
are favored in such conditions. As Pianka (1974) also pointed out, a low
expectation of finding prey, or a high mean search time per item, demands
generalization. The constantly shifting physical parameters typical of
most Apure drainage aquatic habitats probably make it difficult for fish
to concentrate on a given prey species, whose numbers are fluctuating in
conjunction with the physical parameters (Saunders & Lewis 1988b), for
the amount of time required to develop specializations. Hence, most
Apure drainage fishes are omnivores and feeding opportunists.
le 3. Number of
Apure
drainage
characiform
specie
ly) that exhibit
life h
istory strategies K, rl
and r2
K
rl
r2
Total
Anostomidae
-
4
8
12
Characidae
-
50
39
89
Characidiidae
-
11
-
11
Chilodontidae
-
1
-
1
Ctenoluciidae
-
-
1
1
Curimatidae
-
-
8
8
Cynodontidae
-
-
3
3
Erythrinidae
-
-
3
3
Gasteropelecidae
-
-
1
1
Hemiodontidae
-
-
2
2
Lebiasinidae
-
2
1
3
Parodontidae
-
-
2
2
Prochilodontidae
-
-
2
2
TOTALS
0
68
70
138
Characiforms are the most speciose and usually comprise the great¬
est biomass of all Apure drainage fishes in many biotopes. Why is this

839
incredible diversity so great, and how do they sustain such high biomass?
Many authors have addressed the question of high diversity in tropical
fish communities (Roberts 1972, Pianka 1974, Lowe-McConnel1 1975,
Welcomme 1979, Machado Allison 1987). Pianka (1974) provided one of the
best summaries of the various theories in his list of 10 hypothetical
mechanisms of diversity:
1. Evolutionary time. The ancient age of the tropical regions in
terms of evolutionary time allows species to evolve. For example, the
glaciers in North American periodically decimate fishes, the survivors
have not yet reached maximum diversity. This does not happen in the
lowland tropics, and so more time has been available for evolution to
create new species.
2. Ecological time. This hypothesis supposes that the highest tropi¬
cal diversity should occur where fishes have had the longest time to
disperse. Newly created areas (in a geological time-frame) would have
fewer species because of the time necessary for species to disperse
across ecological barriers.
3. Climatic Stability. Stable climate leads to higher diversity.
This idea is being questioned more and more, as more data is available,
and a more precise definition of stability is adopted. Under certain
circumstances this may be true, but in others the very opposite may be
true.
4. Climate predictability. If climate variation is cyclic, it can be
predicted, adapted to and species diversity proceeds.
5. Spatial Heterogeneity. High diversity of niches available to
tropical species allows more speciation. This argument never has been
convincing to me since habitat diversity would seem to be just as rich

840
is many temperate zones where species diversity is much lower than in
the tropics. It is of use to explain local variation in species diver¬
sity, but has little application at the macro-evolutionary level.
6. High productivity, due to more sunlight and warmer temperatures
all year round, accelerates evolution, by allowing the development of
feeding specialists (and eventually, new species). That is, if there is
more food available, you can afford to be "picky."
7. Stability of Primary Production. Plants can buffer climate and
make food available on a more stable basis (or do the opposite). This
hypothesis would suggest that plants make food resources more "conven¬
ient" for year-round consumption in the tropics than in temperate zones
(where winter requires a dormant stage). However, the tropical dry
season would seem to be just as much a problem to tropical plants, and
perhaps even more so. High dry season temperatures without water are
probably more stressful than the low temperatures of winter (when tem¬
perate plants are mostly dormant). Perhaps this explanation is apt for
tropical rain forest regions without a dry season.
8. Competition. This hypothesis holds that natural selection for
competitive ability (K-strategists) is keenest where communities are
diverse. Again, this is assuming a species saturated community. As we
have seen, the Apure drainage is dominated by r-strategists, and in this
respect is more akin to temperate communities that are more governed by
abiotic factors than by biotic interactions.
9. Rarefaction. This hypothesis refers to the continual, density
independent removal of organisms from a community. It suggests that
where communities are not fully saturated with species (such as in the
Apure drainage where climate catastrophically removes most of the fish

841
population each year) more species can coexist. That is, such communi¬
ties are in a sense oversaturated with species, but since losses are due
to chance, it doesn’t really matter.
10. Predation. This argument is similar to 9, in assuming that preda¬
tion, whether selective or random, will always rarefy prey species and
keep them from competing severely with each other. Thus, predators can
suspend the competitive exclusion principal for prey species, by thin¬
ning their populations. Since predation is usually preferential, it is
usually frequency dependent, and would further promote prey diversity.
That is, if predators eat the most common prey item available at any
given time, no one prey species can bloom at another’s expense, and more
can coexist.
In addition to a combination of several of the factors mentioned in
the above hypotheses, the fish communities of the llanos experience
something akin to the "founder effect" of island biogeographers, but it
happens not just once but every year. These fishes reclaim 99% of their
habitat each and every year. The evolutionary implications of this
simple fact are far-reaching. It is as if each year, a small group of
founders arrive to repopulate (in only 8 months for most characiforms) a
recently made "island" (in this case a freshwater sea) virtually unoccu¬
pied by other creatures. We can envision pools (gene and otherwise) of
survivors during the dry season in the mountain springs, in deep river
channels, and in crocodile holes (before they were exterminated by skin
hunters) etc., that shelter a tiny percentage of the total annual bio¬
mass. This seed stock germinates with the first rains, and the recolo¬
nization begins. This yearly reshuffling of the evolutionary deck of
genes has undoubtedly further contributed to the high diversity by

842
reducing the generation time to one year, thus accelerating the evolu¬
tionary clock in a manner reminiscent of the insects.
Although llaneran fishes most likely have an "evolutionary memory"
of the dry season and its consequences, the catastrophic nature of the
mortality suffered means that no individual, no matter what its special¬
izations to survive might be, is certain of survival or even reasonably
positive about it. Thus, generalists are favored, and opportunists
rewarded. Analysis of Table 4 shows that this is precisely the case.
Most species, even those with fairly complex morphological feeding
adaptations, are quite ready to switch to the most abundant food item
available at any given time. Omnivores abound. Of the 137 species of
characiforms studied, 81 were classified as omnivores, 40 as carnivores,
10 as detritivores, and six as herbivores. The high biomass observed in
the Apure drainage (Taphorn & Lilyestrom 1984) is probably partly also a
consequence of the seasonal nature of the environment, and the mandatory
recycling of energy that ensues. Fishes may never exhaust the diverse
food sources that become available as the season advances and the aquatic
habitat continues to expand. Very few food studies of tropical fishes
have taken prey availability into account. This is so because prey items
(usually invertebrates) are hard to identify, and difficult to sample
quantitatively. One study that did take prey availability into account
(Angermeier & Karr 1983) came to the conclusion the distribution (highest
biomass concentration) of several fish species, belonging to various
feeding guilds, were not generally correlated with availabilities of
their major food resources. Their study, however, was carried out in the
dry season, and most species were probably more concerned with simple
survival than optimum food resource utilization. They found that most

843
species were concentrated into deep pools in the Panamanian streams
studied. As more data are available, I believe a pattern will emerge to
show that most fishes in the Apure River drainage are not limited by food
resources. Amazonian blackwater habitats, where the water is nearly
sterile and without dissolved ions of any kind, and pH values drop to
below 4.3, probably provide so little food as to severely restrict fish
populations.
Perhaps because I study them, tropical fishes seem "smarter" than
most temperate fishes. They seem to be more aware of their surroundings
and harder to catch. One behavioral difference I have noticed is the
tremendous drive these fishes have to "get up and go," once it starts to
rain. There seems to be a very strong instinctual urge to move out, and
to do so with haste, and not just in the migratory species but in most
fishes in general. Spurred by rising rivers, most fishes leave their dry
season sanctuaries and move upstream into creeks, and out onto the
flooding savannas. In a short time, every little ditch, pool or pond
that connects into the growing fluvial system as the rains pour down, is
teeming with fishes. They move out to find feeding grounds, to avoid
predators amidst the dense cover provided by the grasses on the flooded
plains, and to spawn. Rapid reproduction is favored in this system.
Fishes are "anxious" to spawn, and many have gonads that ripen (no doubt
at great physiological cost since these fishes are the survivors of the
drought) just before the rains fall. The coporo is such a species. By
spawning in mid-river they suspend their offspring in low-visibility
waters, counting on the rising tide of flood waters to carry them out
onto the plankton-rich floodplains. Dispersal is rapid and widespread.
Many small tetras share the drive to move out and spawn as soon as it

844
rains, which is understandable in light of the different strategies
outlined above.
Most characiform fishes present in the Apure drainage are
characterized by relatively rapid growth. The luxurious aquatic pastures
that form on the flooded savannas are very shortlived and must be quickly
exploited. The ability to attain adult size and reproductive maturity
quickly, and to lay down abundant fat reserves are crucial since these
qualities may determine which of the lucky survivors will make it all the
way through the dry season to successfully spawn. Survival itself is not
sufficient, since individuals must survive in good enough condition to
reproduce almost immediately, before major food resources again become
available after the first rains. Under these conditions, small species
are probably favored (most characiforms are under 60 mm at maturity).
Because predators abound, rapid growth is a must. The faster you can get
bigger, the better. It is a hurry-up, boom and crash population curve.
As we learn more about llaneran fishes, it seems that the rich
species diversity is maintained in part at least by the constant state of
flux in the environment, or in other words by instability.
In a year-long study of the fish communities of the modules of
Apure (the modules are large, 20 x 12 km^ sections of sparsely wooded
savannas that have been surrounded by earthen dikes about two meters
high, they maintain a huge shallow lake that gradually dries up during
the dry season, and provides the moisture necessary for cattle and
pastures, see Taphorn & Lilyestrom 1984 for a more complete account), we
observed that greater stability and isolation from the fluctuating
floodwaters led to lower species diversity. In the creeks and recently
formed modules about 120 species of fishes were found, whereas the

845
older, more stable modules, with large areas that remained permanently
flooded from year to year, were found to have only about 40 species, or
only one-third the number of the younger modules.
Kushlan (1976, 1980) found an opposite effect (greater stability
promoting greater species diversity) in the Florida Everglades, indicat¬
ing some very fundamental differences between the subtropical and tropi¬
cal communities. In Florida, during the unusual stable period, large
predators were able to dominate the community that would otherwise nor¬
mally consist of small omnivores.
In the Apure modules, large predators "dominated" in both biomass
totals as well as the actual number of species present, but the large
predators were the among the best adapted to fluctuations in the climate,
and hence the most likely to survive the dry season. I put the word
"dominated" in quotes because one must bear in mind that the image drawn
of a fish community by sampling with seines a few times during the year
only gives us a snapshot of the community structure at the time the
sample was taken, and this from microhabitat sampled. On the floodplain,
most dry season pools are full of large voracious predators and a few
tetras. The trophic webs begin in the spring on the flooded savanna when
millions of tiny fishes are present, most of them either herbivorous
(feeding on epibenthic or epiphytic [on submerged terrestrial grasses]
diatoms and algae) or omnivorous (feeding on both algae and micro¬
crustaceans). These millions of tiny fishes are the real base of the
trophic pyramid. Most of them perish in the first few hours or days of
their lives, and heavy predation continues throughout the year. Thus, the
relative percentage of the carnivores vs the herbivores (or the more
common omnivores) will continually fluctuate as the season advances.

846
Table 4. Classification of Apure drainage Characiforms by:
Life History Strategy (K, rl, r2);
Diet (C = Carnivore, H = Herbivore, 0 = Omnivore, D = Detritivore);
Water Type (B = Black, W = White, C = Clear);
Abundance (R = Rare, U = Uncommon, C = Common, A = Abundant);
Distribution (A = both mountains and llanos, B = llanos, blackwater
habitats, L = llanos, widespread, M = mountains and/or piedmont,
R = llanos, large rivers.
Anostomidae
• (12
species)
Species Strategy
Diet
Water
Abun.
Di st
Abramites hypselonotus
rl
H
W
U
R
Anostomus ternetzi
rl
0
B,C
U
B
Leporellus vittatus
r2
0
W,C
U
M
Leporinus cf granti
r2
H
B,C
R
B
Leporinus friderici
r2
0
W,C,B
C
L
Leporinus sp.
r2
0
W,C, B
C
L
Leporinus sp. "aguaro"
r2
0
B
R
B
Leporinus striatus
r2
0
W,C
C
M
Leporinus yophorus
r2
0
W
U
L
Pseudanos grácil is
rl
0
B
R
B
Pseudanos irinae
rl
0
B,C
U
B
Schizodon isognathus
r2
H
W,C, B
C
L
Characidae (88
species)
Species Strategy
Diet
Water
Abun.
Dist
Acestrocephalus cf boehlkei
rl
C
W
U
R
Acestrorhynchus falcirostris
rl
C
B
R
B
Acestrorhynchus sp "pico largo"
rl
C
B
R
B
Acestrorhynchus microlepis
rl
C
W,C,B
C
B
Aphyocharax alburnus
rl
0
W,C,B
A
A
Astyanax bimaculatus
r2
0
W,C,B
A
A
Astyanax integer
r2
0
W,C,B
C
A
Astyanax metae
r2
0
W,C
C
M
Astyanax polylepis
r2
0
B,C,
C
B
Astyanax superbus
r2
0
W,C
U
M
Astyanax venezuelae
r2
0
W,C
u
M
Brycon bicolor
r2
0
B, C
R
B
Brycon whitei
r2
0
W,C,B
U
M
Bryconamericus sp.
rl
0
W
C
L
Bryconamericus beta
rl
0
W
C
A
Bryconamericus deuterodonoides
rl
0
W,C
C
M
Bryconops affinis
rl
C
B
R
B
Bryconops melanurus
rl
C
B
U
B
Catoprion mentó
r2
C
B
U
B
Charax gibbosus
r2
C
W,C,B
C
L
Cheirodon pulcher
rl
0
W,C, B
A
A
Cheirodontops geayi
r2
C
W
C
A
Colossoma macropomum
r2
H
W,C
C
L
Corynopoma riisei
rl
C
W,C
c
M
Creagrutus cf beni
rl
0
W,C
c
M
Creagrutus bolivari
rl
0
W,C
c
A

847
Table 4. (continued).
Characidae (continued)
Species Strategy Diet Water Abun. Dist.
Creagrutus n. sp.
Ctenobrycon spilurus
Cynopotamus bipunctatus
Engraulisoma cf taeniatum
Galeocharax sp.
Gephyrocharax valenciae
Gnathocharax steindachneri
Gymnocorymbus thayeri
Hemibrycon metae
Hemigrammus anal is
Hemigrammus barrigonae
Hemigrammus cf elegans
Hemigrammus marginatus
Hemigrammus micropterus
Hemigrammus microstomus
Hemigrammus cf mimus
Hemigrammus newboldi
Hemigrammus cf rhodostomus
Hemigrammus cf schmardae
Hemigrammus sp. "arriba"
Hemigrammus stictus
Heterocharax macrolepis
Hyphessobrycon bentosi
Hyphessobrycon metae
Iguanodectes spilurus
Markiana geayi
Megalamphodus cf axelrodi
Metynnis argenteus
Metynnis hypsauchen
Metynnis luna
Microschemobrycon casiquiare
Moenkhausia chrysargyrea
Moenkhausia copei
Moenkhausia dichroura
Moenkhausia lepidura complex
Myleus cf pacu
Myleus rubripinnis
Mylossoma aureum
Mylossoma duriventris
Paragoniates alburnus
Parapristella georgiae
Phenacogaster megalostictus
Piaractus brachypomus
Poptella orbicularis
Pristobrycon striolatus
Pygocentrus caribe
Pygopristis denticulatus
Roeboides affinis
Roeboides dayi
Salminus hilarii
rl
0
w,c
C
M
rl
0
W,C,B
A
L
r2
C
w
C
L
rl
C
w,c
C
A
r2
C
w
U
R
rl
C
W,C, B
A
A
rl
C
B
R
B
rl
0
B,C
C
B
r2
0
W,C
C
M
rl
0
B
u
B
rl
0
B,C
c
B
rl
0
B
u
B
rl
0
W,C,B
c
A
rl
0
B
c
B
rl
0
B
u
B
rl
0
B
R
B
rl
0
B
U
B
rl
0
B
R
B
rl
0
B
U
B
rl
0
W,C, B
C
L
rl
0
B
R
B
rl
c
B
R
B
rl
0
B
U
B
rl
0
B
C
B
rl
0
B
U
B
r2
0
W,C,B
C
L
rl
0
W,C
U
L
r2
0
B
U
B
r2
0
B
R
B
r2
0
B
U
B
rl
0
B
U
B
rl
0
B
U
B
rl
0
B
C
B
rl
0
W,C, B
C
L
rl
0
W,C, B
C
L
r2
0
B
R
B
r2
0
B
U
B
r2
H
W,C,B
R
L
r2
H
W,C,B
C
L
r2
C
W
C
A
rl
C
B
U
B
rl
C
B
U
B
r2
0
W,C, B
C
L
rl
0
W,C,B
C
L
r2
c
B
U
B
r2
c
W,C,B
C
L
r2
0
B
R
B
rl
c
W,C,B
C
L
rl
c
W,C,B
A
L
r2
c
W,C
U
M

Table 4. (continued).
Characidae (continued)
Species
Strategy
Diet
Water
Abun.
Dist
Serrabrycon magoi
rl
C
B
R
B
Serrasalmus altuvei
r2
C
B
U
B
Serrasalmus medinai
r2
C
W,C, B
C
L
Serrasalmus elongatus
r2
C
B
U
B
Serrasalmus irritans
r2
C
W,C, B
C
L
Serrasalmus rhombeus
r2
C
W,C,B
C
L
Tetragonopterus argenteus
r2
0
W,C,B
C
L
Triportheus albus
r2
0
W
R
R
Triportheus angulatus
r2
0
W,C,B
C
L
Triportheus sp. "cola roj
a" r2
0
W,C, B
C
L
Xenagoniates bondi
rl
C
W,C,B
C
L
Unidentified
rl
0
W
R
L
Characidiidae
(11
species)
Species
Strategy
Diet
Water
Abun.
Dist
Ammocryptocharax elegans
rl
0
B
R
B
Characidium cf zebra
rl
0
B,C,
C
L
Characidium sp. D
rl
0
B
U
B
Characidium chupa
rl
0
W
R
M
Characidium sp. K
rl
0
W,C
R
L
Characidium sp. J
rl
0
B
R
B
Characidium cf catenatum
rl
0
W,C,B
A
A
Characidium sp. G
rl
0
W,C,B
A
L
Characidium voladorita
rl
0
W
C
M
Elachocharax junki
rl
0
B
R
B
Elachocharax pulcher
rl
0
B
R
B
Chilodontidae (1
species)
Species
Strategy
Diet
Water
Abun.
Dist
Chilodus punctatus
rl
0
B
U
B
Ctenoluciidae (1
species)
Species
Strategy
Diet
Water
Abun.
Dist
Boulengerella lucia
r2 '
C
B,C
R
B
Curimatidae
(8 species)
Species
Strategy
Diet
Water
Abun.
Dist
Steindachnerina sp.
r2
D
W,C,B
C
L
Steindachnerina argéntea
r2
D
W,C, B
A
L
Curimata cerasina
r2
D
W
C
L
Cyphocharax spilurus
r2
D
B,C
C
B
Curimatella bolivarensis
r2
D
B,C
R
B
Curimatella immaculata
r2
D
B,C
C
B
Potamorhina altamazonica
r2
D
W
R
R
Psectrogaster ciliatus
r2
D
B
U
B

Table 4. (continued).
Cynodontidae (3 species)
Species
Strategy
Diet
Water
Abun.
Dist
Hydrolycus scomberoides
r2
C
W,C,B
C
L
Rhaphiodon gibbus
r2
C
W,C,B
U
L
Rhaphiodon vulpinus
r2
C
W,C, B
C
L
Erythrinidae
(3 species)
Species
Strategy
Diet
Water
Abun.
Dist
Hoplerythrinus unitaeniatus r2
C
W,C,B
C
L
Hoplias macrophthalmus
r2
C
B
R
B
Hoplias malabaricus
r2
C
W,C,B
A
L
Gasteropelecidae (1
species)
Species
Strategy
Diet
Water
Abun.
Di st
Thoracocharax stellatus
r2 '
C
W,C,B
A
L
Hemiodontidae (2 species)
Species
Strategy
Diet
Water
Abun.
Dist
Hemiodopsis argenteus
r2 '
0
B
R
B
Hemiodopsis gracilis
r2
0
B
R
B
Lebiasinidae
(3 species)
Species
Strategy
Diet
Water
Abun.
Dist
Copel la metae
rl
C
B
C
B
Lebiasina erythrinoides
r2
0
W,C
C
M
Pyrrhulina cf lugubris
rl
0
W,C,B
C
A
Parodontidae
(2 species)
Species
Strategy
Diet
Water
Abun.
Dist
Parodon sp. linea
r2 '
0
W,C
C
L
Parodon apolinari
r2
0
W,C
C
M
Prochilodontidae (2 species)
Species
Strategy
Diet
Water
Abun.
Dist.
Prochilodus mariae
r2 '
D
W, C, B
A
A
Semaprochilodus kneri
r2
D
B,C
R
B

850
Distribution within the Apure drainage.
Although each characiform species has specific, perhaps unique
habitat requirements that in most cases are yet to be ascertained, two
basic environmental parameters determine species distribution within the
Apure River drainage: altitude and water type.
The effects of an altitude gradient on fishes has been studied in
many areas (Balón & Stewart 1983). It could just as well be labeled a
slope, temperature or substrate gradient, since all of these parameters
vary with altitude, and the fishes undoubtedly respond to the combined
effect of these and other associated habitat parameters. Examination of
the species distribution maps reveals that of the 137 species present,
16 (12%) are completely restricted to montane habitats (eg. Characidium
voladorita. Parodon apolinari, and Creagrutus cf ben!)> fourteen more
(10%) use montane and well as llaneran habitats, for a total of 30
species of characiforms in the mountains and piedmont regions. There
are finer divisions within these broad categories. For example,
Corvnopoma riisei is found only in the clearer streams of forested
piedmont areas. (See the individual species accounts for more detailed
information). Most species (107 of 137, or 78%) seldom or never venture
out of the llanos (eg. Charax qi bbosus, Moenkhausi a dichroura).
Combined with the 14 species that use both high and lowlands, a total of
121 (88%) use lowland, llaneran habitats. Of the 107 strictly lowland
species collected, five are usually found in or near large rivers, 43
are widespread throughout the llanos, and 60 are found mostly in
blackwater habitats (47 exclusively so). In Table 5, I have listed the
Apure drainage characiforms that are restricted to the mountains and
pi edmont.

851
Table 5. Characiforms restricted to montane and piedmont habitats.
Anostomidae Leporellus vittatus
" Leporinus striatus
Characidae Astyanax metae
" Astyanax superbus
" Astyanax venezuelae
" Brycon whitei-1
" Bryconamericus deuterodonoides
" Corynopoma riisei
" Creagrutus cf beni
" Creagrutus n sp.
" Hemibrycon metae
" Salminus hilarii-2
Characidiidae Characidium chupa
" Characidium voladorita
Lebiasinidae Lebiasina erythrinoides
Parodontidae Parodon apolinari
1. A population of Brycon tentatively identified as Brycon whitei
exists in the lowland, blackwater Aguaro-Guariquito system. Upon fur¬
ther examination that population will probably be determined as taxonom-
ical 1y distinct from the Andean species.
2. Salminus hi 1arii juveniles have been collected from llaneran
streams, but adults have so far only been found in the Andes. The
juveniles may wash down with floodwaters, and then make there way back
upstream after they have grown.
The second basic parameter controlling fish distribution is water
type. About 44% (60 species) of the 137 characiforms present in the
Apure drainage occur only (or mostly) in blackwater habitats (eg.
Copel!a metae, Hemigrammus cf rhodostomus, Anostomus ternetzi). Whether
they are responding directly to some specific water quality parameter
found in such habitats, such as higher water clarity (visibility), lower
pH values, lack of dissolved minerals (low conductivity), or to some
indirect result of these such as increased aquatic macrophyte
populations, different food sources, a sand versus mud substrate is not
known. While certain parameters may effect fish distribution more than

852
others, fishes respond to the ecosystem as a complex whole. In the
ecological section it was shown that aquatic ecosystems in the llanos
fluctuate seasonally. The water quality parameters mentioned above and
many others such as depth, dissolved oxygen concentration, etc., all run
through an annual cycle of change in accord with the wet and dry season.
Thus, the individual dot maps of distribution reflect an average annual
distribution, rather than precise geographical limits for each species.
In fact, some of the sites shown would be completely dry during part of
the year. In spite of these limitations, the maps show a fairly clear
pattern of distribution for most species. In particular the division of
fish distributions in white and blackwater faunas is one of the most
obvious, and is repeated time and again for many species in several
different families. There are three disjunct areas in the Apure
drainage where blackwater habitats exist: northern Apure state in the
Guaritico River system, (Balsa, Caicara and Guaritico Creeks), central
Barinas state in the region south of Ticoporo, and the Aguaro-Guariquito
system of Guárico state. All three areas are characterized by white-
sand soils, and are drained by streams that originate in the llanos
(although in Barinas, Andean streams also pass through the region, but
these tend to come from forested areas and carry fewer sediments that
other Andean headwater streams).
Tallying the data in Table 4 for those species present in each
water type we find 77 (56% of 137 total) species using Whitewater
habitats, 103 (75%) from blackwater streams, and 74 (54%) that were
found in clear water. Since fishing effort was much more intense in the
most common Whitewater habitat, it is at first surprising to find more
species entering blackwater systems. However, if those species that

853
occur only in blackwater habitats (which I have interpreted as the older
fauna, derived from the ancient Guyana Shield fish fauna) are so
specialized as to be restricted to the same, then we would expect more
species that are restricted to black water than to white water. It
follows then, if the whitewater species evolved more recently to exploit
the geologically young (late Pleistocene), and much more productive
alluvial deposits of the llanos, that they should be more generalized
and opportunistic, and able to take advantage of any situation they find
themselves in. Only 15 species are restricted to whitewater habitats.
Looking at the data another way, we find only 15 species found only in
(apparently restricted to) whitewater habitats, none to Clearwater
habitats (probably because there are so few [perhaps none] year-round
Clearwater habitats here), but 47 in blackwater habitats. Species that
use all three water types numbered 43, 13 were found in both black and
Clearwater habitats, and 19 were found in white and Clearwater habitats.
Lumping whitewater with white and Clearwater streams gives us 33
species, but lumping blackwater with black and Clearwater inhabitants
gives us 60, or almost twice as many. These numbers indicate that those
species in blackwater systems are more restricted to the same than those
using whitewater systems.
Of the 60 species that are mostly restricted to blackwater
habitats (Table 6) 49 (82% of all blackwater species) are present in the
Aguaro-Guariquito system, 31 (52%) in northern Apure state, and 14 (23%)
in central Barinas. Although sampling could have been more thorough, a
pattern of decreasing numbers of blackwater species from east to west
emerges. Major blackwater habitats exist in the rivers draining the
Guyana Shield to the southeast, and also in the Capanaparo and Cinaruco

854
Rivers of southern Apure state. The simplest explanation for the
observed pattern is that river distance determines how many species have
been able to invade the three "islands" of suitable blackwater habitat
(in a sea of muddy, Whitewater rivers) from the probable source areas,
the Guyana Shield and the rivers of southern Apure state. Since the
llanos are geologically very young, species from the ancient (600 mil¬
lion years) Guyana Shield blackwater habitats may not yet have had
adequate time to disperse into all suitable habitats.
Zoogeographical Considerations
The Apure drainage forms a small part of the Orinoco Basin. To
understand how the fishes we currently find there may have arrived, as
well as how they have adapted to the conditions they found there, it is
necessary consider the Apure drainage in the larger context of the
Orinoco Basin, and to consider the history of the Orinoco Basin itself.
Today, the Apure river drainage is sandwiched between the Andes Moun¬
tains to the north and west, and the Guyana Shield to the southeast.
Its soils are a mixture of Quaternary sediments that originated in the
Andes and older sandier soils that were washed down from the Guyana
Shield (or perhaps blown in from there during more arid periods) (R.
Schargel, pers. com.). To understand the zoogeographical history of the
Apure drainage, we must first understand that of the Orinoco Basin.
Unfortunately, various factors create difficulties in the zoogeo¬
graphic analysis of any portion of the South American ichthyofauna
(generally poor taxonomic knowledge at both generic and species levels,
incomplete collections coverage, and very few complete phylogenetic
histories for most fishes). Although the Apure River drainage has now
been more thoroughly sampled than probably any other area of comparable

855
Table 6. List of 59 blackwater species, with an indication of their
presence (X) or absence (-) in the three main regions where blackwater
habitats occur in the Apure drainage.
Anostomidae
Anostomus ternetzi
Leporinus cf granti
Leporinus sp. "aguaro"
Pseudanos grácil is
Pseudanos irinae
Barinas Apure Guárico
X X
X
X
X
Characidae
Acestrorhynchus falcirostris
Acestrorhynchus sp. "pico largo
Acestrorhynchus microlepis X
Astyanax polylepis X
Brycon bicolor
Brycon "whitei"
Bryconops affinis
Bryconops melanurus
Catoprion mentó
Gnathocharax steindachneri
Gymnocorymbus thayeri X
Hemigrammus anal is
Hemigrammus barrigonae X
Hemigrammus elegans
Hemigrammus micropterus X
Hemigrammus microstomus
Hemigrammus cf mimus X
Hemigrammus newboldi X
Hemigrammus cf rhodostomus
Hemigrammus cf schmardae
Hemigrammus stictus
Heterocharax macrolepis
Hyphessobrycon bentosi
Hyphessobrycon metae X
Iguanodectes spilurus
Metynnis argenteus
Metynnis hypsauchen
Metynnis luna
Microschemobrycon casiquiare
Moenkhausia chrysargyrea X
Moenkhausia copei X
Myleus sp
Myleus rubripinnis
Parapristella georgiae
Phenacogaster cf megalostictus
Pristobrycon striolatus
Pygopristis denticulatus
Serrabrycon magoi
Serrasalmus altuvei
Serrasalmus elongatus
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X

856
Table 6. (continued).
Barinas
Apure
Guárico
Characidiidae
Ammocryptocharax elegans
-
-
X
Characidium sp. D
-
X
X
Characidium sp. J
-
-
X
Hoplias macrophthalmus
-
-
X
Copel la metae
-
-
X
Chilodontidae
Chilodus punctatus
-
X
-
Hemiodontidae
Hemiodopsis argenteus
-
X
-
Hemiodopsis gracilis
-
-
X
Curimatidae
Cyphocharax spilurus
X
X
X
Curimatella bolivarensis
X
X
-
Curimatella inmaculata
-
X
X
Psectrogaster ciliatus
-
X
-
Prochilodontidae
Semaprochilodus kneri
Barinas
Apure
X
Guárico
total number of
species:
13
30
49
percent:
22
51
83
size in South America, the dearth of collections from other portions of
the Orinoco basin, as well as from neighboring river basins, permit only
tentative zoogeographic conclusions at this time.
Another factor is our incomplete knowledge of the geological and
hydrological history of northern South America, although recent work by
Duque-Caro (1989) permits some preliminary conclusions to be drawn,
which are summarized below. The history of Andean orogeny begins in the
middle Eocene, when intermontane basins began to take shape, and the
emergence of the Cordillera Central divided northern South America into
eastern and western regions. The Guyana Shield and the Cordillera

857
Oriental, the latter of which extend from west to east through Venezuela
out towards Trinidad, had already partially emerged by this time. Major
paleogeographic and paleo-oceanographic events during the middle Miocene
further shaped the drainages in this region, and by the Middle Pliocene
events had proceeded to the point that mountain ranges and river basins
had, to a large degree, attained their present-day configurations.
During the formative stages, before the Andean uplift had proceeded very
far, there is evidence that a major river ran north from southern
Colombia, or perhaps even northern Ecuador, out through the present-day
Magdalena Basin and Panama (the Central American land bridge had not yet
emerged at this time.
Fossil evidence indicates that serrasalmines and other fishes now
extinct west of the Andes were present in the region at that time. As
the Andes rose, this river’s delta migrated eastward, possibly flowing
into the Caribbean first through what is now the Maracaibo Basin, and
then through central Falcon state in Venezuela, and the Uñare drainage
area in eastern Venezuela, to finally reach its present day delta near
Trinidad. This scenario is greatly oversimplified, (eg., parts of the
area were flooded by ocean during part of this time), but in the absence
of more precise geological information it can serve as a working
hypothesis.
The similarities of the characiform fishes found today in the
Apure drainage clearly indicate a closer relationship between the ich¬
thyofaunas east of the Andes than with those west of the Andes.
To get an initial idea of the possible relationships and origins
of the Apure drainage fishes, one can compare the lists that are
available in the literature for neighboring drainages. Eigenmann was

858
always concerned with the origin of fish faunas as well as their
taxonomy and diversity, and published many lists of the fish species
occurring in different drainage areas. In his work on the fishes of
British Guiana (1912) he published such a list comparing the Orinoco
Basin fishes then known (only 89 species!) with the 470 species he
obtained from Guyana (then British Guiana). Using the admittedly
incomplete (taxonomically) list of characiform species for the Apure
River drainage in Venezuela presented here as the starting point, Table
7 was constructed to compare Apure fishes with those reported from
surrounding drainages for which lists are available. Lists for the
Guyana made by Eigenmann (1912), for the Rio Negro in Brazil (Goulding
1988), from the Maracaibo Drainage in Venezuela (Taphorn & Lilyestrom
1984), and from the Magdalena Drainage of northern Colombia (Dahl 1971)
were used. For some records of species not in the original lists were
included, based on more recent collections of my own or reported in the
literature. Although the limitations listed above are serious, a fairly
clear pattern of faunal similarity emerges. The Apure River drainage
shared many more families, genera, and species with the Rio Negro
drainage and Essequibo Basin, than with the Maracaibo or Magdalena
basins on the other side of the Andes.
As can be seen in the summary of the data at the end of Table 7,
of the thirteen families present in the Apure drainage the Chilodonti-
dae, Cynodontidae and Hemiodontidae are absent from both the Magdalena
and Maracaibo basins. Genera provide the most useful information
regarding affinities of the Apure drainage characiform fishes. Of the
77 genera present, only about 20 occur today west of the Andes. More
than 76% of the genera present in the Apure drainage are also found in

859
the Amazonian blackwater tributary, the Rio Negro, and about 64% are
shared with the Essequibo drainage in Guyana. Since specific
identifications are usually tentative, little more than a general trend
can be detected with such an analysis. Probably the most important
information to convey, however, is that only seven or eight species are
common to both sides of the Andes, whereas between 25-30% of the Apure
species are common to Guyana or the Rio Negro.
It seems clear then that the Apure drainage characiform fish fauna
is comprised of both old and new elements. The old elements were proba¬
bly present in the Guyana Shield region long before the Apure drainage
was formed. If a complete species list were available for the upper
Orinoco, I suspect that many more species and genera would be found in
common with the Apure fauna than is true for the Rio Negro. The high
degree of similarity between the Apure characiforms and the Rio Negro
species is due to the large number of blackwater species found in iso¬
lated patches in the former. Those species have probably dispersed from
the Shield into the Apure drainage. The newer element of the Apure
drainage fish fauna consists of species that have evolved there. One
group has specialized in utilizing mountain and piedmont habitats, and
another evolved to take advantage of lowland, whitewater rivers. This
mixture of origins probably accounts in part for the relatively high
species diversity observed.

860
Table 7. Zoogeographical comparison of the Apure Drainage characiform
ichthyofauna. Comparisons are given for the Magdalena Basin of northern
Colombia (Magd.); the Maracaibo Basin of northwestern Venezuela (Mara.); the
Essequibo and Demerara Basins of Guyana (Guya.); and the Rio Negro Drainage
of northern Brazil (Negr.). Taxa in common are indicated by F = Family, G =
Genus and S = Species. Sources are indicated in text. An asterisk indi¬
cates that the taxa was not mentioned in the original source but is included
based on subsequent records.
Anostomidae
Abramites hypselonotus
Anostomus ternetzi
Leporellus vittatus
Leporinus cf granti
Leporinus friderici
Leporinus sp.
Leporinus sp. "aguaro"
Leporinus striatus
Leporinus yophorus
Pseudanos gracilis
Pseudanos irinae
Schizodon isognathus
Magd. Mara.
F F
S
S
Guya.
F
S
G
S
S
Negr.
F
S
S
S*
S
S
G G
S
S
G
Characidae F
Acestrocephalus cf boehlkei G
Acestrorhynchus falcirostris
Acestrorhynchus sp "pico largo"
Acestrorhynchus microlepis
Aphyocharax alburnus
Astyanax bimaculatus S
Astyanax integer
Astyanax metae
Astyanax polylepis
Astyanax superbus
Astyanax venezuelae
Brycon bicolor
Brycon whitei S
Bryconamericus sp. G
Bryconamericus beta
Bryconamericus deuterodonoides
Bryconops affinis
Bryconops melanurus
Catoprion mentó
Charax gibbosus
Cheirodon pulcher G
Cheirodontops geayi
Colossoma macropomum
Corynopoma riisei
Creagrutus cf beni S
Creagrutus bolivari
Creagrutus n. sp.
Ctenobrycon spilurus
Cynopotamus bipunctatus
Engraulisoma cf taeniatum
F
G
G
S
G
S
G
F
S
S
G
S
S
G
G
S
S
S
S
G
S
S
G
F
G
S
S
S*
G
G
G
S
S
S
G
G

861
Table 7. Continued.
Galeocharax sp.
Gephyrocharax valenciae
Gnathocharax steindachneri
Gymnocorymbus thayeri
Hemibrycon metae
Hemigrammus anal is
Characidae (continued)
Hemigrammus barrigonae
Hemigrammus cf elegans
Hemigrammus marginatus
Hemigrammus micropterus
Hemigrammus microstomus
Hemigrammus cf mimus
Hemigrammus newboldi
Hemigrammus cf rhodostomus
Hemigrammus cf schmardae
Hemigrammus sp. "arriba"
Hemigrammus stictus
Heterocharax macrolepis
Hyphessobrycon bentosi
Hyphessobrycon metae
Iguanodectes spilurus
Markiana geayi
Megalamphodus cf axelrodi
Metynnis argenteus
Metynnis hypsauchen
Metynnis luna
Microschemobrycon casiquiare
Moenkhausia chrysargyrea
Moenkhausia copei
Moenkhausia dichroura
Moenkhausia lepidura complex
Myleus cf pacu
Myleus rubripinnis
Mylossoma aureum
Mylossoma duriventris
Paragoniates alburnus
ParapriStella georgiae
Phenacogaster cf megalostictus
Piaractus brachypomus
Poptella orbicularis
Pristobrycon striolatus
Pygocentrus caribe
Pygopristis denticulatus
Roeboides affinis
Roeboides dayi
Salminus hilarii
Serrabrycon magoi
Serrasalmus altuvei
Serrasalmus elongatus
Serrasalmus irritans
Serrasalmus medinai
Magd. Mara. Guya. Negr.
G G
S
G G
S
S
S*

862
Table 7. Continued.
Characidae (continued)
Serrasalmus rhombeus
Tetragonopterus argenteus
Triportheus albus
Triportheus angulatus
Triportheus sp. "cola roja"
Xenagoniates bondi
Characidiidae
Ammocryptocharax elegans
Characidium cf catenatum
Characidium chupa
Characidium sp. D
Characidium sp. G
Characidium sp. K
Characidium sp. J
Characidium voladorita
Characidium cf zebra
Elachocharax junki
Elachocharax pulcher
Chi 1odontidae
Chilodus punctatus
Ctenoluciidae
Boulengerella lucia
Curimatidae
Curimata cerasina
Curimatella bolivarensis
Curimatella immaculata
Cyphocharax spilurus
Potamorhina altamazonica
Psectrogaster ciliatus
Steindachnerina argéntea
Steindachnerina sp.
Cynodontidae
Hydrolycus scomberoides
Rhaphiodon gibbus
Rhaphiodon vulpinus
Magd. Mara.
Guya. Negr.
S S*
S G
G
S
Erythrinidae
Hoplerythrinus unitaeniatus
Hoplias macrophthalmus
Hopli as malabaricus
F F
S S
F F
S
S S*
S S*
Gasteropelecidae
Thoracocharax stellatus
F F F F

863
Table 7. Continued.
Magd.
Mara.
Guya.
Negr.
Hemiodontidae
F
F
Hemiodopsis argenteus
Hemiodopsis gracilis
G
G*
Lebiasinidae
F
F
F
F
Copel la metae
G
Lebiasina erythrinoides
G
S
G
G*
Pyrrhulina cf lugubris
G
G
Parodontidae
F
F
F
F
Parodon sp. linea
Parodon apolinari
G
G
G
G*
Prochilodontidae
F
F
F
F
Prochilodus mariae
G
G
G
G
Semaprochilodus kneri
present
G*
in
Number
Shared with
Apure
Magd.
Mara
Guya.
Negr.
Families
13
10
10
13
13
Genera
77
20
19
49
59
Species
138
8
7
37
39

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South America. Stanford Ich. Bull., 7(4):109-113.
. 1960b. Further notes on the relationships and classification
of the South American characid fishes of the subfamily Gasteropeleci-
nae. Stanford Ich. Bull., 7(4):217-239.
. 1960b. The phylogenetic relationships of Triportheus, a genus
of South American fishes. Stanford Ich. Bull., 7(4):239-244.
. 1962. The osteology of Brvcon meeki. a generalized characid
fish, with an osteological definition of the family. Stanford Ich.
Bull., 8(1)1-77.
. 1964. Osteology and relationships of South American characid
fishes of subfamilies Lebiasininae and Erythrininae with special
reference to the subtribe Nannostomina. Proc. U. S. Nat. Mus.,
116:127-170.
. 1986. A new species of Elachocharax (Teleostei: Characidae)
from the Rio Negro region of Venezuela and Brazil. Proc. Biol. Soc.
Washington, 99(4):739-747.
Weitzman, S. & J. Cobb. 1975. A revision of the South American fishes
of the genus Nannostomus Giinther (Family Lebiasinidae). Smithsonian
Cont. Zool., 186:1-36.
Weitzman, S. & J. Géry. 1981. The relationships of the South American
pygmy characoid fishes of the genus Elachocharax, with a redescrip¬
tion of El achocharax .iunki (Teleostei: Characidae). Proc. Biol. Soc.
Washington, 93(4):887-913.
Weitzman, S. & R. Kanazawa. 1976. Ammocryptocharax eleoans. a new genus
and species of riffle-inhabiting characoid fish (Teleostei: Characi¬
dae) from South America. Proc. Biol. Soc. Washington, 89(26):325-
346.
. 1977. A new species of pygmy characoid fish from the Rio Negro
and Rio Amazonas, South america (Teleostei: Characidae). Proc. Biol.
Soc. Washington, 90(1):149-160.
. 1978. The South American fish genus Elachocharax Myers with a
description of a new species (Teleostei: Characidae). Proc. Biol.
Soc. Washington, 91(1):158-183.
Winemiller, K. & D. Taphorn. 1989. La evolución de las estrategias de
vida en los peces de los llanos occidentales de Venezuela. Biollania 6.
Winterbottom, R. 1980. Systematics, osteology and phylogenetic rela¬
tionships of fishes of the ostariophysan subfamily Anostominae
(Characoidei, Anostomidae). Life Sciences Contribution: 123. Royal
Ontario Museum.
Zinck, A. 1977. RIOS DE VENEZUELA. Cuadernos Lagoven. Caracas.

APPENDIX
NUMBER OF SPECIMENS EXAMINED
Lots Specimens
Anostomidae
Abramites hypselonotus 14 28
Anostomus ternetzi 19 69
Leporellus vittatus 16 49
Leporinus cf granti 4 6
Leporinus friderici 64 230
Leporinus sp. 25 81
Leporinus sp. "aguaro" 3 3
Leporinus striatus 35 109
Leporinus yophorus 11 84
Pseudanos gracilis 5 8
Pseudanos irinae 6 17
Schizodon isognathus 92 296
SUBTOTALS: 294 980
Characidae
Acestrocephalus cf boehlkei 13 80
Acestrorhynchus falcirostris 1 1
Acestrorhynchus sp "pico largo" 2 37
Acestrorhynchus microlepis 46 334
Aphyocharax alburnus 323 6992
Astyanax bimaculatus 319 6704
Astyanax integer 106 691
Astyanax metae 70 1002
Astyanax polylepis 41 642
Astyanax superbus 23 278
Astyanax venezuelae 10 105
Brycon bicolor 3 3
Brycon whitei 24 42
Bryconamericus sp. 77 3096
Bryconamericus beta 140 2838
Bryconamericus deuterodonoides 117 5187
Bryconops affinis 1 11
Bryconops melanurus 19 256
Catoprion mentó 24 187
Charax gibbosus 144 985
Cheirodon pulcher 431 21330
Cheirodontops geayi 101 1705
Colossoma macropomum 0 0
Corynopoma riisei 28 362
Creagrutus cf beni 87 1315
887

888
APPENDIX, (continued).
Lots Specimens
Creagrutus bolivari 71 759
Creagrutus n. sp. 33 575
Ctenobrycon spilurus 355 16691
Cynopotamus bipunctatus 31 220
Engraulisoma cf taeniatum 31 129
Galeocharax sp. 22 306
Gephyrocharax valenciae 211 5636
Gnathocharax steindachneri 2 2
Gymnocorymbus thayeri 157 11035
Hemibrycon metae 87 1190
Hemigrammus anal is 11 147
Hemigrammus barrigonae 57 1373
Hemigrammus cf elegans 6 50
Hemigrammus marginatus 4 125
Hemigrammus micropterus 184 3767
Hemigrammus microstomus 37 2096
Hemigrammus cf mimus 16 594
Hemigrammus newboldi 23 252
Hemigrammus cf rhodostomus 4 116
Hemigrammus cf schmardae 16 607
Hemigrammus sp. "arriba" 149 4733
Hemigrammus stictus 1 1
Heterocharax macrolepis 2 5
Hyphessobrycon bentosi 21 112
Hyphessobrycon metae 28 348
Iguanodectes spilurus 20 348
Markiana geayi 124 878
Megalamphodus cf axelrodi 17 83
Metynnis argenteus 24 178
Metynnis hypsauchen 3 8
Metynnis luna 15 89
Microschemobrycon casiquiare 6 41
Moenkhausia chrysargyrea 6 38
Moenkhausia copei 42 1378
Moenkhausia dichroura 118 2210
Moenkhausia lepidura complex 44 722
Myleus cf pacu 1 1
Myleus rubripinnis 10 13
Mylossoma aureum 0 0
Mylossoma duriventris 49 203
Paragoniates alburnus 61 222
ParapriStella georgiae 8 462
Phenacogaster megalostictus 17 183
Piaractus brachypomus 15 31
Poptella orbicularis 108 3662
Pristobrycon striolatus 12 40
Pygocentrus caribe 174 1136
Pygopristis denticulatus 1 4
Roeboides affinis 111 826
Roeboides dayi 279 8067
Salminus hilarii 15 26
Serrabrycon magoi 2 7

APPENDIX, (continued).
Lots
Specimen
Serrasalmus altuvei
6
14
Serrasalmus medinai
68
216
Serrasalmus elongatus
17
51
Serrasalmus irritans
142
744
Serrasalmus rhombeus
61
136
Tetragonopterus argenteus
35
178
Triportheus albus
5
181
Triportheus angulatus
39
105
Triportheus sp. "cola roja"
107
1045
Xenagoniates bondi
87
574
Unidentified
1
219
SUBTOTALS:
5559
129371
Characidiidae
Ammocryptocharax elegans
1
1
Characidium cf zebra
26
191
Characidium sp. D
8
17
Characidium chupa
7
9
Characidium sp. K
4
36
Characidium sp. J
6
24
Characidium cf catenatum
82
389
Characidium sp. G
83
464
Characidium voladorita
46
302
Elachocharax junki
1
7
Elachocharax pulcher
2
8
SUBTOTALS:
266
1448
Chi 1odontidae
Chilodus punctatus
24
156
Ctenoluciidae
Boulengerella lucia
10
11
Curimatidae
Steindachnerina sp.
27
182
Steindachnerina argéntea
314
4229
Curimata cerasina
69
253
Cyphocharax spilurus
78
2495
Curimatella bolivarensis
7
32
Curimatella immaculata
103
2809
Potamorhina altamazonica
12
20
Psectrogaster ciliatus
4
16
SUBTOTALS:
614
10036

890
APPENDIX, (continued).
Cynodontidae
Lots
Specimens
Hydrolycus scomberoides
14
80
Rhaphiodon gibbus
8
22
Rhaphiodon vulpinus
12
66
SUBTOTALS:
34
168
Erythrinidae
Hoplerythrinus unitaeniatus
38
120
Hoplias macrophthalmus
2
2
Hop!ias malabaricus
252
649
SUBTOTALS:
292
771
Gasteropelecidae
Thoracocharax stellatus
161
2777
Hemiodontidae
Hemiodopsis argenteus
2
2
Hemiodopsis gracilis
2
31
SUBTOTALS:
4
33
Lebiasinidae
Copel!a metae
9
64
Lebiasina erythrinoides
32
407
Pyrrhulina cf lugubris
157
2198
SUBTOTALS:
198
2669
Parodontidae
Parodon sp. "linea"
24
344
Parodon apolinari
44
229
SUBTOTALS:
68
573
Prochilodontidae
Prochilodus mariae
114
348
Semaprochilodus kneri
4
6
SUBTOTALS:
118
354
TOTAL
7642
149347

BIOGRAPHICAL SKETCH
Mr. Donald Charles Taphorn was born in Belleville, Illinois, on the
eighth of September in 1951 to Mrs. Shirley Jean Taphorn nee Baechle and
Mr. Donald Clemens Taphorn, as the first of five children. His interest
in fishes began with the gift of a goldfish from his grandmother, and as
an aquarist the hobby soon mushroomed from one goldfish bowl to a basement
full of tropical fish aquaria.
After attending Catholic grade school and high school in Belleville,
he enrolled as a biology major at Southern Illinois University at Edwards-
ville. There he met ichthyologist Dr. Jamie E. Thomerson, who hired him
to work in the aquarium room, and later introduced him to the tropics and
annual killyfish. In 1972, he graduated from SIUE, and was awarded an
Outstanding Biology Senior Award. That same year, he moved to Gaines¬
ville, Florida, to enroll in the graduate program of the Department of
Zoology at the University of Florida. His major professor was Dr. Carter
Gilbert, curator of fishes at the Florida State Museum. After finishing
the necessary course work, he began work on a revision of the cyprinodon-
tid genera Austrofundulus and Rachovia, the topic of his masters degree,
which was completed in 1976.
In 1976 he accepted an offer of employment from DISCA (Department of
Environmental Contamination Research) of the Venezuelan Ministry of the
Environment. From 1976-1977 he worked with Mr. Craig Lilyestrom on an
inventory and food-habit study of the fishes of the Lake Maracaibo
891

892
Basin, in search of fishes that might be useful in the biological control
of mosquito larvae.
From 1977-1978, he worked at the University of Zulia in Maracaibo,
teaching fish systematics in the Department of Hydrobiology. Since Janu¬
ary, 1979 he has been a member of the faculty of the Universidad Nacional
Experimental de los Llanos Occidentales Ezequiel Zamora (UNELLEZ), a
federal university in Guanare, state of Portuguesa, as a Professor of
Vertebrate Zoology and Aquatic Resource Management in the Environmental
Engineering Department, and as Curator of Fishes and Director of the
Museum of Zoology. In 1989 he was appointed as coordinator of the Gradu¬
ate Program in Wildlife and Fisheries Management being developed jointly
by UNELLEZ and the International Office of the U. S. Fish and Wildlife
Division.
His research is dedicated to the taxonomy and biology of freshwater
Venezuelan fishes. Current research topics include an inventory of the
other (non-characiform) fishes present in the Apure River drainage, and a
review of the fishes of the family Rivulidae in Venezuela being carried
out jointly with Dr. J. E. Thomerson. He has published over thirty-five
scientific papers on neotropical freshwater fishes.

I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
AtiidJAlkfr
Carter R. Gilberyc, Chairman
Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy. ,
Mr,M
Horst Schwassmann
Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
Frank Nordlie
Professor of Zoology
I certify that I have read
conforms to acceptable standards
adequate, in scope and quality,
Doctor of Philosophy.
this study and that in my opinion it
of scholarly presentation and is fully
as a dissertation for the degree of
/
Richard Kiltie
Associate Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
Nigel Smith
Professor of Geography
This dissertation was submitted to the Graduate Faculty of the
Department of Zoology in the College of Arts and Sciences and to the
Graduate School and was accepted as partial fulfillment of the require¬
ments for the degree of Doctor of Philosophy.
May, 1990
Dean, Graduate School

m
£).
UNIVERSITY OF FLORIDA
3 1262 07332 005 2





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