Sei sulla pagina 1di 127

Cover: Hyphessobrycon serpae. Photo by H.J. Richter.

Quotations of reasonable length in

scientific, non-com-

charge and

mercial pUblications are allowable without without permission from the publisher.

ISBN 0-87666-458-3

~~iy

Printed

in the U.S.A.

@Copyrlght

1977 by T.F.H. Publlcatlons, Inc. Ltd.

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2

PREFACE

This book, devoted to the most diversified group' of American and African fresh-water fishes, has two goals. First, to provide the general reader, who is an aquarist more often than not, with as many good illustra- tions of aquarium type characoid fishes as possible. This kind of pictorial guide will enable him, it is hoped, to identify at first sight a good portion of the aquarium species. At the same time, the non-specialized reader will get an idea of the position of the species in the classification(genus, family and so on) and, if he wants, can refer to the text for more systematic and, occasionally, biological data, using the index of Latin names at the end of the volume. The second aim of the book is to give to zoologists as well as to advanced aquarists a scientific account of our present knowledge of the group, which has not been reviewed as a whole for balf a century. This account has been made as complete and reliable as possible, with a minimum of words and a bibliography reduced to a few recent works, but it is by no means to be taken as a true revision of the suborder. In order to classify and to describe all, or aImost all, known species and enable curators of museum cóllections and other interested specialists to identify them with reasonable accuracy, dichotomous tables, also called identification keys or dichotomous keys, are presented along with a discussion of the various species and their particular characteristics. The author is aware that such a system causes a kind of psychological block to certain persons. These readers are asked to content themselves with the illustrations and general statements regarding the groups, which should be quite sufficient in most cases. Others, it is hoped, will make the simple effort to enter the field of "yes or no" reasoning, which is basically the computer's, as well as our own brain's, way of thinking. To master the use of a key, which only takes a few minutes, the principIe is as follows: one is asked, step by step (a, b, c, etc.) a question to which the answer is either yes or no. If the organism to be identified has such or such a feature (answer yes), one goes to the next step. If not (answer no), one goes to the corres- ponding branch of the altemative (represented by aa, bb, cc, etc.), which is more or less remote in the key, until the identification is achieved.

L!:cb

c.

Po.<;;t

;. >:c.,

d:

'i"".

.':':-;1

:-~'P~!1

Py -Danlel

69,000 - Marwu

- Amo.o

- B,ulJ

3

-----------.

II

Actually, this will probably not always be the case, and the reader must

be prepared for some false leads. The impossibility of identifying a particular species may be due to the foJlowing: certain keys are surely imperfect, suffering from some defect wbich can only be detected after several tests; certain exceptions t9 the presence or absence of a character may cause difficulties, although they should be mentioned in the key when known; and the whole group is still imperfectly known and it may be, cbiefty if the species is small and comes from some remote area, that the unidentifiable

specimen has never

been scientifically described (a so-called new species).

The characoids now number more tPan 1200 species (inc1uding 200 or so Mrican ones), and, based on past experience, in every large collecting

sample 5-10% of the species can be expected to be new.

4

ACKNOWLEDGMENTS

The

author

wishes to extend to the book:

contributed

bis thanks

to the persons

who have gready

Dr. Warren

Burgess,

who had the difficult task of making

the author's

language readable by English-speaking people, and the people whose

remarkable photographs provide the indispensable illustrations, principally

Dr. H. R. Axelrod is dedicated.

Schultz, in whose memory this book

and the late Harald

5

-~

I

('

Table of Contents

Preface.

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Acknowledgments

 

5

Introduction

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9

Part 1:

The African FamUies.

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Key to the African Families

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. 17

Family

Hepsetidae

(African pike-like characoids)

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Family

Alestidae (African tetras).

 

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Subfamily

Alestinae

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Subfamily

Hydrocyninae

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Family Citharinidae

(citharinids)

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Subfamily Citharininae

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Subfamily

Distichodinae

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.58

Subfamily

Ichthyborinae

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.82

Part 2:

The American

Characoids

other than the Characidae.

 

.95

 

Key to the American Families

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.95

Family

Family

Family

Erythrinidae

Ctenoluciidae

Crenuchidae.

(trahiras)

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. (South American pike-characoids). .

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. 106

Family

Characidiidae

(South

American

"darters")

 

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Subfamily Characidiinae.

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SubfamilyElachocharacinae

 

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Subfamily

Geisleriinae

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. Family Lebiasinidae (pencilfishesand pyrrhulinins)

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. 122

.123

Subfamily Lebiasininae

 

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Subfamily Pyrrhulininae Family Anostomidae (headstanders and leporins)

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Subfamily Leporellinae

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.151

 

Subfamily

 

.151

Family Hemiodidae (hemiodids and their relatives)

 

.190

Subfamily Hemiodinae

.190

Subfamily Parodontinae

 

202

Family Curimatidae (curimatas).

 

.210

Subfamily Chilodinae

 

211

Subfamily Prochilodinae

 

.214

Subfamily Curimatinae

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7

 

Subfamily Anodinae

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. (hatchetfishes) (pacus, silver dollars and piranhas) .

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Family Gasteropelecidae Family Serrasalmidae

Subfamily

Subfamily

Myleinae

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Subfamily

Catoprioninae.

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:. 294

Part

3: The American

Tetras

and Characins

(Family Characidae)

 

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Subfamily

Agoniatinae.

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298

Subfamily

Rhaphiodontinae

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Subfamily

Characinae

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. Subfamily Clupeacharacinae

Subfamily

Bryconinae

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346

Subfamilies

Paragoniatinae

and Aphyocharacinae

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SubfamilyGlandulocaudinae

 

.351

Subfamily

Stethaprioninae

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366

Subfamily

Tetragonopterinae

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Subfamily Rhoadsiinae

538

Subfamily Cheirodontinae

and allied genera

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543

Undescribed

or unidentified

species

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.609

Glossary of technical

terms

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.611

Bibliography

I: Africa

621

Bibliography

11:

South America

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.642

Index

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.647

8

INTRODUCTION

Onee upon a time, an Mrican fresh-water fish decided to beeome a eharaeoid.

This happened long ago, in the Mesozoie (70-175 millions of years?) when eonditions were favorable. Volcanism was intense, working profound modifications of the earth. Our fish lived iJ) a mythical land, called Gond- wanaIand, formed by the united continents of Mrica and South America. They were soon to be separated, owing to a gigantie break which had just started, forming a ehain of great lakes. A similar process is actualIy in action in 'Mrica today; the new break starts with the Red Sea and is prolonged by the Rift VaIley forming the Great Lakes of Africa (Tanganyika, Malawi, Victoria, ete.). Sueh lakes are the most suitable plaees to alIow what is calIed explosive evolution to oecur because they offer the greatest ecologica1opportunities to the species. These species are able to oecupy a multitude of empty spots sueh as rocks, sand, grass-beds, bottom, surfaee, and so on, or potentiaI so-calIed eeological niehes. Another possible cause of rapid and active speeiation in Great Lakes, as eontrasted with the "normal" evolution in

adjaeent rivers, may be the laek of important

Hydrocynus or Lates in certain Mrican lakes) whieh, when present, restrain the diversification of the species.

fish predators

(sueh as

The ambitious fish mentioned at the beginning of the story (which is not

a fairy-tale but a hypothesis based on eertain welI established facts), was probably a species resembling what we now calI a gonorynchiform. That smalI group of partIy fresh-water fishes, mostly Mrican, with Kneria, Cromeria and others, seems indeed to have some "pre-ostariophysine" features. An ostariophysine fish (eharaeoids were apparently the first ostario-

physans, although certain authors believe that the catfishes are more ancient)

is eharacterized by at least two features: a chain of ossic1es,calIed Weber's

ossic1es,uniting the gas-bladder with the internal ear and permitting the hearing of high-frequency sounds; and a chemical alarm system, telling the sehooling fish that one of its eongeners has been attacked by a predator. These two adaptations are supposed to be useful to fishes living in schools in fresh-water biotopes (which is the case in inost characoids and cyprinoids).

9

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Recent investigations have shown that most, if not ali, of the Kneria-like

61

of these structures. They have specialized

cervical vertebrae (the first three) as if they were in the process of differen- tiating ossicles (which is achieved in the characoids), and they also have a typical fright reaction if a wounded fish of their kind (or a cyprinid) is introduced into a school.

recent fishes do possess rudiments

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With ali the above cited conditions brought together, our Kneria-like hypothetical procharacoid was roughly in the same position as a cichlid was,

rather recently, in a great Mrican lake, that occupied alI potential (empty

or not completely exploited) niches and gave rise to the famous, extremely

diversified, recent fauna ofthe Great Lakes. It is likely that the procharacoid

literally underwent explosive radiation with such a spread of adaptations

that there are no other such examples in the vertebrates except for the Australian marsupiaIs.

This evolution was probably possible thanks to the general organization,

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with several generalized features, which characterizes a characoid. In this

group, there fs usually an adiposefin and, almost always, weli developedscales.

Teeth are alwayspresent, at leastin the post-Iarva. There is no rostralbone,no

is direct/y articulated with the Weberian ossicles are

complete,with the tripusfixed on the body 01 the third vertebra by means 01a

Q) vibrating lamelia, in contrast to ali other ostariophysans. Once the continents were separated, the above defined characoids evolved,

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as preethmoid, and the prevomer is not toothed and

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as

-

the palatine head and lateral ethmoid. Final/y,

for their own part, on each side of the newly-formed Atlantic OceanoIt was

very probably impossible for these strictly so-called primary fresh-water

fishes to survive in marine water, except for short periods of time or short

distances.

It is believed (but not by ali zoologists) that in Mrica they soon gave rise

to the cyprinids, of which the primitive stock could have been Barilius-like

fishes (?), which in turn radiated and became ardent competitors of their

own parents (the descendants of the procharacoids) in the very same

biotopes. These barbs emigrated to Asia and North America by the

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lasciatus reached Texas by a different route, the southem

Central America). In America, the characoids gave rise to the highly specialized gymnotoids after the separation of the continents. Gymnotoids never developed in Mrica,

where their niches are occupied by knife-fishes and mormyroids.

Neither the gymnotoids nor the catfishes (of unknown origin), specialized very early, were such competitors as were the barbs in Mrica. The American (Neotropical) characoids were thus free to diversify in almost ali possible directions. In addition, the drift of the continents had caused the

one through

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upheaval of the Andes, as if the westem coast of South America had abutted against a static block in its course to the west. This in turn caused the formation of the Amazon basin, the greatest of the world, which offered the best opportunities to a young, opportunistic group of fishes. These two reasons explain why the Neotropica1 characoid fauna repre- sents, in number of species, about 80% of the total, and the African ("Ethiopian") fauna consequently only 20%, with 11 and 3 families res-

pectively, according to certain modem authors.

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On each side of the Atlantic Ocean, similar ecological pressures deter-

mined a parallel evolution. Certain evolutionary convelgences are worth mentioning:

The African pike-characoidHepsetushas its counterpart in the

American Acestrorhynchus.

The large African alestids (except Hydrocynus) strongly parallel the

American Bryconinae, whereas the small alestids are scarcely discemable

from the numerous American tetras.

Hydrocynus, a modification of an alestid toward predation, has dose similarities with the Neotropical Hoplias or Erythrinus.

Most of the Citharinidae are convergent with certain Hemiodidae and

Curimatidae (two rather dosely related American families), whereas

some others, very small and bottom-living fishes (Nannocharaxand allies),

are strikingly similar to the Neotropical Characidiidae.

1n the above cited examples, only knowledge of the origin, or the study of anatomical details, can tell the non-specialist whether he is dealing with an African or an American line, which diverged from each other so many millions of years ago. Hepsetus, the most primitive characoid according to T. Roberts (forming the family Hepsetidae), has an accessory ectoptery- goid bone and a canal into the supraoperde. These characters are not shared by any Neotropical characoid, even the Ctenoluciidae which are apparently the dosest group. The citharinids have ctenoid scales(except for one genus), numerous ventral rays, pharyngeal "bags" and other features which do not

exist (at least together) in American families. Finally, the alestids differ

ãi

"-

CD

c:

CD

ti)

ai

-

o

fi)

:J

-

f!!

ai

Q.

Q.

ai

c:

ai

".:; from their dosest American relatives, the Characidae, in having no rhino-

.8 CD

sphenoid, a bony tube endosing the olfactory nerve, a toothless, paddle-like

maxilla, and other characters. 1t has been pointed out above that the evolution of the characoids, chiefty of the Neotropical ones, resembles that of the more recent cichlids from certain African lakes. Both groups, each with a different time scale however, entered new biotopes and exploited all of their possibilities. 1ndeed, mos! of the adaptations enumerated by Fryer and Iles (1972) conceming the African cichlids are found in the characoids. They practice commensalism, mimicry,shoaling (also called schooling),migration, burrow-

13

-1

ing in sand, and so on. They have developed three means of communication, namely sounds, chemicals, and recognition marks (mostly on humeral region .and caudal pedunc1e, as welI as on dorsal and adipose fins). They have, in certain cases, elaborated or evolved breeding habits such as the care of the young as practiced by the piranhas. Finally, they have developed an array offeeding habits, which is largely comparable to that ofthe cichlids, inc1uding scale eating or fin biting, as well as other adaptations which are almost unique among teleosts, such as flight (hatchet-fishes) or group- predation (piranhas).

Owing to the

millions of years of evolution at the

disposal 0f the

characoids, these adaptations ocçurred more than once and in different phylogenetic lines. This resulted in an extremely confusing complexity of groups. It is often difficult to telI from a character shared by different forms the role of true phylogeny (common origin) and of convergency (different

origin). That is how a generalized body-shape, like that of a salmon or a trout for example, is to be seen at the same time in Brycon, Salminus and Astyanax, which originated from three different evolutionary lines (although in the same family). Evolution of a predatory existence, that is predation mostly on fishes, appeared at least eight times in completely different lines: Hep~'etus,Hydro- cynus and certain Citharinidae in Africa, Ctenoluciuset aI., Erythrinus et aI., Acestrorhynchus et aI. and several other Characinae, not forgetting the piranhas (Serrasalmidae) in South America. The plant-eaters belong to at least three families, namely Citharinidae, Anostomidae and Curimatidae. But some members of other groups, for example Deuterodon in the Tetragonopterinae, and several others, also developed a herbivorous habitoThe same is true for the limnivorous species, found in the Citharinidae as welI as in the Curimatidae, the "Aufwuchs"- eaters, which nibble food from rocks or falIen trees (two or three families), the plankton-feeders (at least two families), and the specialized forros referred to above. Scale-eaters appear in two families, Characidae and Serrasalmidae, and the fin-biters also in two families, Citharinidae and (probably) Serrasalmidae. Several species are adapted to life in river torrents. They belong to three families, Citharinidae (with Nannocharaxet al.), Characidiidae (with Chara- cidium et aI.) and Hemiodidae (with Parodon et al.). A ventral keel appeared in several subfamilies of the Characidae (Trip- ortheus, Gilbertolus,Gnathocharax, Pseudocorynopomaand so on), and also in certain Curimatidae, as welI as in the Serrasalmidae, culminating, in the Gasteropelecidae, in a complex of adaptations permitting flight a few inches above the water surface. FinalIy, sexual dimorphism is evident in the Characidae as welI as the

Alestidae, Serrasalmidae, Crenuchidae and even in one curimatid genus, whereas care of the young appeared in at least three lines, certain Erythrini- dae and Serrasalmidae, and certain Pyrrhulininae in the family Lebiasinidae. The above enumeration of their intricate adaptations helps to explain why the systematics of the characoids has long been, and stilI is, a difficult task. They were recognized as a homogeneous group only in 1844 by two German ichthyologists, MülIer and Troschel. The Viennese Kner and his student Steindachner (between 1858 and 1915) folIowed by accurately des- cribing a number of species without paying much attention to their c1assi- fication. It remained for one of the best ichthyologists of alI times, Carl H. Eigenmann, to undertake, in the U.S., a natural c1assificationof the South American groups. His principal monographs were issued between 1910 and

was folIowed by his student, George S. Myers, the still very active

father of the "characologists," who in turn handed the torch down to his

1927. He

own

students,

J. Bõhlke,

S. Weitzman

and several others,

forming

a chain

of interest which is presently stilI growing. Recentlya series of anatomical studies by S. Weitzman, T. Roberts, and Others helped specify the systematic position of several critica1 groups, which led to the recognition of many families within the suborder (a division which could never have been accepted in Eigenmann's time) and revealed the importance, as welI as the complexity, of the characoids, presently considered as a key-group among the teleosts.

into several families (16)

was done by S. Weitzman, in Greenwood et aI. (1966). The arrangement of the families in the present book takes into account that c1assification,as modified by Géry (1972), with a few alterations from the 1972 order.

As compared with Weitzman's folIows:

The first authoritative division ofthecharacoids

c1assification, the

present

one

is

as

S. WEITZMAN in GREENWOOD

et a!.

(16 families)

(1) CHARACIDAE

(2) ERYTHRINIDAE

(3) CTENOLUCIIDAE

(4) HEPSETIDAE

(5) CYNODONTIDAE

(6) LEBIASINIDAE

(7) PARODONTIDAE

(8) GASTEROPELECIDAE (9) PROCHILODONTIDAE

ARRANGEMENT

(14 families) (2) ALESTIDAE; (6) CRENUCHIDAE; (7) CHARACIDIIDAE; (13) SERRASAL- MIDAE; (14) CHARACIDAE (4) ERYTHRINIDAE (5) CTENOLUCIIDAE (1) HEPSETIDAE

PRESENT

RHAPHIODONTINAE,

a subfamily

of the

CHARACIDAE

 

(8) LEBIASINIDAE

PARODONTINAE,

a subfamily

ofthe

HEMIODIDAE (12) GASTEROPELECIDAE

PROCHILODINAE,

a subfamily

of the

CURIMA

TIDAE

(10) CURIMATIDAE

(11) ANOSTOMIDAE (12) HEMIODONTIDAE

(13) CHILODONTIDAE

(11) CURIMATIDAE (including PROCHILODINAE, CHILODINAE and ANODINAE) (9) ANOSTOMIDAE

(10) HEMIODIDAE

PARODONTINAE)

CHILODINAE,

CURIMA

(including

a subfamily

of the

TIDAE

--

Part

The African

I

Families

-

(14) DISTICHODONTIDAE

 

DISTICHODINAE,

a subfamily

of the

 

CITHARINIDAE

 

(15) CITHARINIDAE

 

(3) CITHARINIDAE

(including

 

DISTICHODINAE

and

KEY TO THE AFRICAN

FAMILIES*

ICHTHYBORINAE)

(16) ICHTHYBORIDAE

 

ICHTHYBORINAE,

a subfamily

of the

 

CITHARINIDAE.

 

a. Scalescycloid; teeth usually well developed, not movable; joint between two sides

accessory ectopterygoid bone;

a large supraopercle bearing a lateral line

The evolutionary importance of the group will be conspicuous, it is hoped,

of mandible with some interlocking device, movable; no pharyngeal "bags". b. Predaceous fishes armed with strong canines; maxilla entirely toothed; an

in the body of the book, which

will deal successively

with the African and

canal

HEPSETIDAE

American characoids,

ending,

in

a third

part,

with the most successful

bb. Omnivorous

fishes

usually

with

incisiform

and

molariform

teeth, rarely

Neotropical

group,

the Characidae.

 

piscivorous and with canines (as in the subfamily Hydrocyninae,

 

genus);

maxilla

never

toothed;

no

accessory

ectopterygoid

with a single bone; no

supraopercle

ALESTIDAE

 

(2 subfamilies)

 

aa. Scales ctenoid

(except

in Citharinus,

which

is a bream-like

fish very different

from

Hepsetus

or

from

an

alestid);

teeth

usually

small,

often

movable;

joint

between

two sides of mandible

simple, sometimes ankylosed;

pharyngeal

"bags"

present

CITHARINIDAE

 

(3 subfamilies)

 

Family Hepsetidae

 

African Pike-like Characoids

Hepsetusodoe,formerly called Sarcodacesodoe,is the single representative of the family. Its external resemblancewith anAcestrorhynchus,by convergent evolution, is striking. However, an examination of the finer structures of its skull reveals a number of differences. Hepsetus, according to Roberts, is more closely related to another South American group, the Ctenoluciidae, but with many more primitive features (accessory ectopterygoid bone, flattened cranial roof, large canal-bearing supraopercle, and so on) which make it probably the most primitive characoid. The African pike lives in almost alI rivers of the tropical area, with the exception of the Nile basin. It is a fish-predator, welI armed with strong, caniniform teeth, similar to those of Hoplias, Acestrorhynchus and the like. The maxillary bone is

* This key, like ali other keys in the book, is for identification purposes only. It is not intended to reflect relationships.

16

17

entirely toothed as in most predators. According to the author's own experience in Mrica, it is a good game-fish, behaving like a European pike, putting up a good fight. It reaches 30 em in length and is edible. Its color is brownish with indistinct bars on the body and small spots on the fins; its adipose fin is bright orange and black, apparently forming a recognition

signal.

Svensson, as well as Johnels, has reported that Hepsetus spawns somewhat

in the manner

spawning pair and the eggs (which are afterwards guarded by both parents or one parent only) are deposited in it.

of the

anabantids.

A nest of floating foam

is made

by the

Family Alestidae

African Tetras

The Alestidae are the Mrican counterparts of the Characidae, and there is little doubt that the two groups have the same originoOne example will show how similar they are. A new genus and species (Pellegrininahetero- lepis) was once described as a new Mrican form dose to Alestesand Brycinus, because the specimen studied was thought to come from "some part ofWest Mrica." A more critical evaluation revealed afterward that the label on the bottle was incorrect and that the fish was a well-known South American Chalceus. Despite the general resemblance between the two groups (which led

several ichthyologists to recognize on1y one family from both sides of

the

Atlantic Ocean), the Alestidae have the following characteristics in con-

trast to those of the Characidae (at least those characids which most resemble the alestids, namely the Bryconinae): no rhinosphenoid (a small

bone in front of the orbitosphenoid, present in Brycon); olfactory nerve endosed in a bony tube formed by the orbitosphenoid and the lateral

when present, very low, the number of scales between

lateralline and ventral fin Dever exceeding 3 (usually 2). In the Alestinae, the maxillary bone (more or less covered by the first infraorbitals) is tooth- less, usually paddle-like with a rounded contour and a thin pedicel (like a music note). The premaxillary bone has a characteristic pedicel-like posterior process extending to the maxilla, which is rather firmly united to it; the inner premaxillary teeth are invariably four in number (apparently a primitive number, also to be found in the American Hemibryconini), and molariform (except in the small species), usually with the crushing face excavated, the posterior border rounded, multi-cuspidate, and the anterior one with two prominent cusps or points. On the mandible there is usually a pair of small conical teeth behind the main series.

ethmoid; lateralline,

18

.-

In the Hydrocyninae, which are ichthyophagous predators derived from some Alestes, the powerful jaws are armed with a single series of very acute caniniform teeth; it is believed that these teeth correspond to the outer series of the alestins. Another pecu1iarityofthis group is the type ofmiddle joint between the two parts of the mandible. It is a "bolted hinge," whereas that of the alestins is said to be "a fluted tenon." Otherwise, as concems the general organization, a Hydrocynus does not ditfer' essentially from an the Alestes. subfamiliallevel: Accordingly, the family Alestidae will be split into two groups at

a. Teeth in 2 rows on the upper jaw (three rows in one genus), the inner row usually composed of molariform teethj omnivorous species

ALESTINAE

aa. Teeth in 1 row on both jaws, caniniformj ichthyophagouS species

HYDROCYNINAE

SUBFAMIL Y ALESTINAE

African

Tetras

Quite artificially, one may divide the alestins into two groups, Alestini

and Petersini, which could be called tribes. The Petersini look like the young of the Alestini and constitute what are properly called the Mrican tetras.

They ditfer from the Alestini not on1y by their

correlation with the reduction of the lateralline), but also in other "regres- sive" characters. The inner teeth of the upper jaw are no longer excavated

small size (frequently in

and molariform, but have retained a single, pluricuspidate cutting edge, and the inner mandibulary conical teeth are often lacking. Some alestins, the so-called longipinnis-group,form a SOftoftransition between the two groups, evident. the species being small and the molariform structure of their teeth scarcely

Tribe Alestini The presentauthor recognizes3 generadiagnosedby the followingkey:

a. An adipose lid covering at least part of eye; always a fontanel; scales with radiating "radii" which are never anastomosed.

b. Premaxillary teeth triserial; 3 scales between lateral line and ventral fin;

dorsal fin well in front of midbody,

2.8-3.5instandardlength)

filamentous

in adult male j body

deep

Bryconaethiops

(depth

bb. Premaxillary teeth biserialj It-2

3.25-5instandardlength)

and ventral finj

dorsal fin at midbody or even behind, not filamentous; body elongate (depth

scales between latcralline

A/estes

lia. No adipose lidj

usually no fontanel, except in a few species (B.

B. /ongipinnis,B. chapen',B. opisthotaenia)j

anastomosed

"radii"

of

the

Brycinus

scales

leuciscus,

usually

.

.

"

~9.000_

,.,

"":>

Py.

Vaniel

8,

,.

C I.C''- po~rltll.;',- .P \

Manaus-.

Amazona.-

I

19

I

GENUS

BRYCQNAETHIOPS

This genus includes a few species whieh ehange eonsiderably during growth, eertain of the adults being seareely identifiable with the young. By the time they reaeh 200 mm in length they are usually very deep, very dark (sometimes almost blaek), and the filaments of the dorsal fin of the males reaeh the caudal finoThe group is at onee reeognized by the 3 series of teeth on the upper jaw (it is somewhat like modified Alestes, just as Creagrutus,in the American tetras, looks like a modified Hemibrycon),but is urgently in need of a revision at the species leveI. The two or three speeies, B. microstoma,B. boulengeriand perhaps B. macrops(whieh, if valid, is mueh rarer), are mostly from the Congo basin (including Ubanghi) although some species are found in Cameroon and Gaboon.

Bryconaethíops

sp.

Photo by Dr. J. Gery.

GENUS

ALESTES

A few species, namely Alestes dentex, A. baremose,A. macrophthalmus,

A. liebrechtsiiand A. stuhlmanni, have rather reeently been separated from

the bulk of the Alestes-like species on the basis of anatomieal details sueh as

adipose lid, fontanels, ete. They are usually large (up to 45 em), elongate speeies inhabiting large rivers. Some are of eeonomie importanee. Aleste.~ dentex and A. baremose,from the Nile, Niger, Tehad, Senegal, ete., are sibling species, being separable only by the number of gill-rakers.

Alestes macrophthalmusis a southem form from the Congo and Gaboon, A. liebrechtsiiis restricted to the C~ngo basin, and the small A. stuhlmanni seems to be an endemic form from East Mrica. Exeept for some red eolor on the caudal fin in eertain forms, species of Alestes are usually uniform silvery, without eharacteristie markings.

A/estes baremose and A. líebrechtsíí (photos by Dr. J. Gery); A. dentex (after Boulenger, 1909); A. macrophtha/mus, holotype (after Boulenger, 1909).

GENUS BRYCINUS

This genus is an aggregate of species in which one may recognize five groups. Some of these could, perhaps, be ranked as subgenera, although overlapping of characters is frequent; the others are quite artificial

assemblages.

The

Brycinus

macrolepidotus -group

(1), which

contains

the

type species of the genus Brycinus, B. macrolepidotus,and a few other

species, is characterized by the large size (up to 50em), straight dorsal profile, dorsal fin far behind mid-body, and overhanging snout. The other

four groups,

which have dissimilar characters,

may be recognized as

foUows:

a. Interorbital usually less than 3 times in length of head; caudal peduncle high (its depth 1.0-1.5 in its length). b. 16-28 gill-rakers on lower arch (fontanel always absent; scales usually with

anastomosed radii) c. Dorsal fin slightlyin front of mid-body cc. Dorsal fin at mid-body or slightly behind

.

.

.

(2) Brycinus nurse-imberi-group

B. nurse

B. imberi-group(6-7spp.)

bb. 11-18 gill-rakers on lower arch (fontanel present or absent; scales with or without anastomosed radiO.

d. Bodydepth 3.5-3.66in standardlength; 4 sca1esbetweendorsalfin and

lateral line; sca1es without anastomosed radii; no fontanel; dorsal fin

usually behind mid-body

(3) B. rutilus-group(2 spp. ?)

dd. Body depth 2.3-3.5 in standard length; 5-6 sca1esbetween dorsal fin and

sca1eswith anastomosed radii; sometimes a fontanel; dorsal

lateralline;

fin usually at mid-body or in front of mid-body (4) B.longipinnis-group (8-9 spp.) aa. Interorbital 3.0-3.5 times in length of head; depth of caudal peduncle 1.5-2.0 in

its length (fontanel always absent; scales without anastomosed radii)

.

.

.

(5)

B.

humilis-group

(4 spp. ?)

(1) The Brycinus macro/epidotus-group The large species belonging to the nominal genus Brycinus have a dis- tinctive shape, the straight snout (which continues the long, also straight, predorsalline) overhanging the mandible. The smallest of the six induded species, B. brevis (225 mm) from Nigeria and the Gold Coast, is also the deepest (body depth 2.75 to 3.0 in standard length), with few lateral line sca1es(21-23), and is a uniform color, Brycinuscarmesinusis very dose to it. The four other species are more elongate (body depth 1.5 to 4.0 in the standard length). Two of them, B. grandisquamisfrom the Congo basin and B. batesiifrom South Cameroon, attain 26 em in length and can be easily recognized thanks to a conspicuous spot on the end of the caudal pedunde. They have, respectively, 23-27 and 28-29 lateral line scales, and 13-14 versus 15-16 anal rays.

22

"""

.'

~~

4

jj-

:"

Brycinus sp., juvenlle. Photo by G. Timmerman.

Brycinus macrolepidotus, the type species ofthe genus, inhabits a large part of Mrica from the Nile to the Congo. It usually has a broad dark longi-

tudinal

scales (22-26 sca1es in B. macro-

lepidotus versus

differing only in the number

bando Brycinus rhodopleura is its represen~tive

of lateralline

in Lake Tanganyika,

28-29 in B. rhodopleura).They reach, respectively, 42 and

30 em in length.

(2) The Brycinus nurse-imberi-group

used much

nowadays) and the Brycinus imberi-group (with B. imberi, B. jacksoni, B. affinis, B. taeniurus, B. kingsleyae and B. bimaculatus), are medium-sized (100-230 mm) alestins with the dorsal fin at mid-body or thereabouts and 16-28 gill-rakers (usually less than 18 in the Other groups).

Brycinus

nurse (type

species of the genus

BracJzyalestes, not

Brycinus nurse is the only species in this group to have the dorsal fin slightly in advance ofmid-body, and it is also the largest (230 mm); it has the same geographical distribution as B. macrolepidotus, pIus Lakes Victoria and

Rudolf.

The caudal fin and part of the ventral and anaI fins are usually red;

humeral

and caudal

spots are present.

Species ofthe B. imberi-group strictly speaking have the dorsal fin inserted at mid-body or slightly behind. B. kingsleyae and B. taeniurus from South

Cameroon, Gaboon and Chiloango are immediately recognizable by the dark lateral band beginning behind the dorsal fin leveI and ending on the middle caudal rays. Brycinus kingsleyae has a humeral spot (Iacking in B. taeniurus) and a shorter anaI fin (12-13 branched rays versus 15-17).

23

B. imberi (of which curtus, bequaerti and jacksoni are possibly synonyms) from Cameroon to Kasai, B. bimaculatus from the upper Congo, and B. affinis, an endemic species from East Mrica, are mostly distinguished by their color pattern. B. imberi usually has a conspicuous humeral spot and a broad caudal spot ending on the caudal rays; B. bimaculatus has the same caudal spot but it is restricted to the caudal pedunde and also has a lateral spot much farther back (at the leveI of the end of the dorsal fin); B. affinis

has the same humeral

(3) The Brycinus

spot as B. imberi, but has

rutilus-group

no caudal spot at all.

This small group differs from the preceding one mostly in having fewer gilI-rakers and from the next one by the structUre of the scales and the more

.

B;ycinus bimaculatus, hoJotype (after Boulenger, 1899) and B. nurse (photo by Dr. J. Gery).

24

r

elongate body. B. rutilus is a large form (up to 32 em) with the fins some- times bright vermilion (hence its name). lt inhabits the same regions in West Africa as B. macrolepidotus,but is nowhere captUred with it (a vicarious

known species

species). B. stolatus from the Congo is a small (?), poorly without color marks.

(4) The Brycinus longipinnis-group

This quite unnatUral group forms a sort of transition with the small alestins called tribe Petersini. Most of the species are small. They some- times have a fontanel and, at least when young, their inner premaxillary teeth are more compressed than in typical Brycinus. However, a dose examination reveals a pair of smalI "shoulders" in front of the largest teeth, which is almost the only character which distinguishes them from the next tribe.

B. leuciscus,from the Senegal and Niger basins (where it is calIed tineni

and is of economic importance), has long been confounded with the young of B. nurse, but it has yelIow fins instead of red. Another species, more recently described, is also very dose. This is B. dagetifrom the Tchad basin, which differs from B. nurse in being matUre at half the size of the latter

species, as welI as by the number of gilI-rakers. It seems to be the sibling of

B. nurse and might welI be dassified together with B. leuciscus,within the

B. nurse-group. The B.longipinnis-group

strictly speaking is in need ofrevision

(like other

Brycinus groups).

The

folIowing key will aid in distinguishing

the species:

a. Not more than 30 lateralline

scales (18-21 branched

tous in male; caudal band present)

.

.

.

anal rays; dorsal fin filamen-

B.longipinnis,B. chaperi

and an undescribed forro from the Ivory Coast (West Africa, plus Equatorial Africa for B. chaperi

on1y ?)

aa. 32-37 lateralline

scales.

b. 16-21 branched anal rays.

c. An elongate caudal spot; 16-17 branched anal rays; 33 lateralline

scales

B. sadleri (endemic species

from Lake Victoria)

cc. Caudal spot broad or absent; 17-21 branched anal rays; 33-37 lateralline scales. d. A broad caudal spot; dorsal rays filamentous in male, marked with a black

bar;

33-35 lateral

line

scales

.

.

dd. Body and fins plain; 371ateralline scales

.

B. intermedius(South

Cameroon)

. B.tessmanni(South

Cameroon)

bb. 24-26 branched anal rays (body and fins plain; 32 lateralline

scales)

B. tholloni (Congo basin)

25

B. longipinnisand B. chaperi,probably sympatric in West Mrica, are the best known alestins among aquarists. The two species are difficu1tto teU

apart, especially when one has to dea1 with young individuaIs. As a role, B. chaperi has slight1ymore lateralline sca1esthan B. longipinnis (28-30 versus 26-27), as welI as a longer snout, shorter maxilIary bone, broader

interorbital and deeper caudal pedunde.

behind farther (versus back than slight1yin that of front B. longipinnis, of mid-body). being at mid-body or slight1y

Its dorsal fin is also inserted

(5) The Brycinus

The

four

species.

humi/is-group

B.

humilis,

B.

abelt~ B.

lateralis

and

B.

opisthotaenia

are

also very dose to certain species of Micralestes.They are smalI (maximum size 150mm), with few scales (24-34) and few branched anal rays (12-17). Brycinus opisthotaenia,from South Cameroon and Gaboon, is the largest species and has retained many Brycinus macrolepidotuscharacters. It. is remarkable in having contact tuberdes (similar to those on the snout and

(photo 8rycinus by humilis, Dr. J. Gery). hOlotype (after Boulenger, 1909) and 8. rutilus

26

head of mature male barbs and species of Labeo) over the whole body in

both sexes. Nothing is known about the significance of these tuberdes (calIed nuptial tuberdes or pearl organs in the barbs). Like B. opisthotaenia,

B. lateralishas a black lateral band (after preservation) but no humeral spot.

It inhabits Katanga and Angola up to the Zambezi River, and differs by having more scales (30-33 versus 24-27 lateral line scales in B. opisthotaenia).

B. humilis from Angola and B. abeli from Ubanghi have a horizonta11y

elongate caudal spot; they differ in the same way as the above, having respectively 30 and 24-26 lateral line sca1es.

Tribe

Petersini

5mall

African

Tetras

This group of about 60 species has recent1y been revised by PolI. He

some proposed by Hoedeman in

195.6which are questionable. It looks as if the group is oversplit, which seems evident when one compares the Petersini with their American counter-

13 genera with about 80

species. However, the evolutionary conditions were not exact1ythe same,

owing to the absence in South America, for example, of the barbs, which

Here on1ya part of the 16 genera

not described at

(actually 18 induding Ladigesia and Virilia, which were

admitted 16 genera as valid, induding

part the Hemibryconini. The latter group has 12 or

compete severely with the Mrican tetras.

the time of PolI's revision) will be accepted as diagnosed by the following key:

aoTeeth multicuspidate; two premaxillary series.

b. Predorsalline

scaled; no outward-directed teetho

c. A pair of conical or (rarely) cuspidate teeth behind the main mandibulary series.

d. Teeth usually with more than 3 cUSPSo e. More than 4 outer premaxillary teeth total (lateralline usually complete)

(I) Micralestes

eeo4 outer premaxillary teeth, 2 on each side (lateral line complete or

incomplete)

o o o (2) Phenacogrammus

(This genus may be split into 3 or 4 groups) dd. Teeth tricuspidate (lateralline incomplete; 4 outer premaxillary teeth)

(3)

Tricuspidalestes

cc. Conical teeth behind the main mandibular series usually absento f. Scales of fiank and abdomen of equal size; 4 or 6 outer premaxillary teeth (2-3 on each side); 8-11 mandibular teeth. g. Posterior fontanel present; mandible not prominent (lateral line

completeor incomplete)

o(4) Hemigrammopetersius (This genus may be split into 4 or 5 groups of species)

"

gg. Posterior fontanel absent, at least in adults; mandible very pro-

minent (lateralline complete)

(5) Petersius

ff. Scales of flank much larger than those of abdomen; 8 outer pre-

maxiUary teeth; 16-20 mandibular

teeth

bb. Predorsalline

. (6) Arnoldichthys

naked; teeth partly outward-directed

. (7) Ladigesia

aa. Teeth conical, in one series (lateralline incomplete).

h. Scales normal; mouth cIupeoid

(8)

Clupeocharax

hh. Bodynaked,withoutscales;mouth normal

The Petersini

are thus

composed

.

.

.

(9)

of 6 monotypic

Lepidarchus

genera which

are very

well 'Characrerized (Tricuspidalestes, Petersius, Arnoldichthys, Ladigesia, Clupeocharax and Lepidarchus) and of three polytypic genera, two of which are quite complicated and perhaps not monophyletic.

(1)

GENUS MICRALESTES

Micralestes is the lar,gest genus of the Petersini as well as the most generalized.All the speciesare of small size (maximum length 100or 110mm for the type species, Micralesteshumilis),are moderately elongate, usually have a longitudinal dark band, and are very similar to each other. According to Poll, they can be distinguished as follows:

t",

M/era/estes

aeut/dens.

Photo by Dr. Herbert R. Axelrod.

a. Pair of small teeth behind main mandibulary series conical, without cusps; no more than 7-9 cusps in the broadest teeth. b. Usually less than 29 lateralline scales. c. No black longitudinal band; body depth 2.6-2.9 in standard length

. M. holargyreus(Congo basin) cc. A bIack longitudinal band, more or less complete; body depth 2.9-4 in standard length. d. Eye 2.6-3.3 in length of head (adult); more than 50 mm in standard length. e. 22-24 Iateralline scales.

f. Longitudinal band beginning at dorsallevel; head 3.7-3.9 in standard

length; 15-16branched anal rays

M. stormsi(Congobasin; very difficultto separate from M. humilis)

ff. Longitudinal

band

complete;

16-18branchedanalrays

head 3.95-4.1 in

standard

M. vittatus(Lake

length;

Tanganyika) ee. 25-29 lateralline scales (longitudinal band complete). g. Lateralline complete. h. Length of head 3.4-3.8 in standard length. i. Scales ofIateraI line with bIack spot at base; 25-27 lateralline

scales

.

.

. M.humilis(Tchad,Congo

and Zambezi Rivers)

i i. Scales of lateralline

not marked;

24-25 Iateralline

scales .

hh. Length

. M. voltae (upper Volta basin; difficult to teU from M. elongatus)

of head 3.75-4.1

in standard

length.

j. 18-19 branched anal rays; tip of dorsal fin black (25-26

lateralline

scales)

j j. 16-17 branched

M. occidentalis(WestMrica)

anal rays; tip of dorsal fin hyaline

. M. sardina (upper Congo

basin) gg. Lateral line usuaIIy not quite complete, lacking a few pores on caudal peduncIe; 16-19 branched anal rays; length of head 3.4-4 in standard length (species of West Mrica). k. Tip of dorsal fin black; body depth 3.45-3.95 in standard

length

M.elongatus(upperVolta)

kk. Tip of dorsal fin hyaline; body depth 2.8-3.3 in standard

length

M.comoensis(IvoryCoast)

dd. Eye 2.25-2.5 in length of head (adult); less than 50 mm in standard

length

M./odori(Congo)

bb. 29-33 lateralline band complete).

scales (body depth 3.5-4.2 in standard length; longitudinal

1. 16-19 branched marked.

anal rays; scales of lateral line not

m. Caudal peduncIe 1.15-1.4 times longer than deep

. M. lualabae(Congobasin)

mm. Caudal peduncIe 1.5-1.75 times longer than deep M. argyrotaenia(Angola)

ll. 19-21 h.:r-mchedana1 rays; sca1es of lateral line with black spot at base (caudal peduncle as deep as long)

.

M.

congicus(Congo basin)

aa. Pair of small teeth behind main mandibulary series with 2-4 cusps; up to 12 cusps in the broadest teeth (longitudinal band complete; tip of dorsal fin black; 14-16

branched ana1rays and 26-29lateral line sca1es)

M. acutidens(fromNigeria

to Angolaand Zambezi basin; WestMrica?)

(2)

GENUS

PHENACOGRAMMUS

This

composite

incomplete

lateral

genus

inc1udes

the

Micralestes-like

species

with

an

line (however

Micralestes elongatus and comoensis some-

times have an incomplete lateralline), plus certain forms which probably come from di1ferent evolutionary groups but are not separable at the generic leveI, their characters showing some overlap. These were considered

as genera

a

lower leveI, as follows:

by Poll.

It seems convenient

to retain

a similar division,

but

at

a. Lateralline

complete.

" ~]

'

'

,

,I

I

fI .

11.,

i~

b. 4t or 5t transverse sca1esbetween predorsal line and lateralline

bb. 6t-7t

. Ph. altus-group (or subgenus Brachypetersius) and lateralline

. Ph. ansorgii-group(or

transverse sca1esbetween predorsalline

subgenus

Nannopetersius)

aa. Lateral line incomplete (or not quite complete in certain individuaIs) (4t-8t

transverse sca1esbetween predorsalline

and lateralline).

c. 15-21 branched anal rays; unpaired fins often filamentous in adult male; body of moderate depth (2.6-4.3 in standard length)

cc. 22-25

usually

branched

anal rays;

deep (body

depth

. Ph. interruptus-group

(Phenacogrammus

typica1)

unpaired

fins unrnodified

lengt